MONILOPHYTA / FERNS s.l.

Roots with a single apical cell, originating from the pericycle, lateral roots from the
endodermis; stem with apical initial(s), plasmodesmatal density in whole SAM 1956[mean]/m2 [lineage-specific mitochondrial network]; stem with hypodermal and
outer-cortical band of fibres [= stereome]; amphiphloic siphonostele +, discontinuities in
stele in t.s. caused by frond gaps; protoxylem restricted to lobes of central xylem strand
[giving a beaded appearance, hence monilophytes], xylem mesarch, tracheids with
scalariform pits, G-type tracheids in protoxylem; phloem fibres rare; stem endodermis
and pericycle +; leaves megaphyllous [ad/abaxial symmetry evolved first, then
determinancy], development acropetal; petiole with multiple leaf traces coming from a
U-shaped bundle; frond veins not anastomosing; sporangia grouped in sori,
sporangium stalk 6< cells across, walls two cells thick, dehiscence by an exothecium,
tapetum amoeboid, spores/sporangium 1000<, white, globose-tetrahedral, with
orbicules, wall development centrifugal, exospore 3-layered, pseudoendospore +;
gametophyte thalloid; antheridium wall 5 cells thick, male gametes with 30-150 cilia,
with numerous plastids and mitochondria; root lateral with respect to the longitudinal
axis of the embryo [plant homorhizic]; x = ca 121, genome size [1C] = ca 14.3 pg
[averages], chloroplast rps4 gene with nine-nucleotide insertion, LSC inversion
from trnG-GCC to trnT-GGU; loss of one group II mitochondrial intron.
Age. Magalln et al. (2013) estimated a crown-group age of around (404-)394.3389.9(-382) m.y. for this clade, ca 360 and ca 364 m.y. are the ages in Schuettpelz and
Pryer (2007) and Y-L. Qiu et al. (2007) respectively, (390.7-)368.5(-354) m.y. in Zhong
et al. (2014b) and around 330 m.y. in Villarreal and Renner (2014); see also P. Soltis et
al. (2002) for suggestions.
Evolution. Divergence & Distribution. Schuettpelz and Pryer (2009, esp. Tables 2, 3
in the Supplement) and Rothfels et al. (2015b: Appendix S4) provide extensive dating of
divergence times in monilophytes, and for more ages of the major monilophyte clades
see Y.-L. Qiu et al. (2007) and Schneider et al. (2004a). Schuettpelz and Pryer (2009)
also list a number of fossil records (see also Rothwell & Stockey 2008).
There have been several radiations of homosporous leptosporangiate ferns, the first
in the Palaeozoic, giving rise to lineages that have since become extinct, and again in the
Jurassic and the Cretaceous (Rothwell & Stockey 2008). General fern diversity decreased
(along with that of the cycads) through the Cretaceous (Wing & Boucher 1998). Ferns
appear to have temporarily dominated at least locally after the end-Cretaceous bolide
impact/Deccan Traps eruptions (Schneider et al. 2004a). The eusporangiate Marattia and
Angiopteris and the leptosporangiate royal and tree ferns are almost living fossils and
show little molecular and even morphological evolution (P. Soltis et al. 2002; Zhong et al
2014b; Rothfels et al. 2015b; Bomfleur et al. 2014a, c.f. Schneider et al. 2015).
Some 98% of living ferns are leptosporangiate ferns, and their main radiation may
have have been after the diversification of the angiosperms in the late Cretaceous-early
Caenozoic; initially it was a largely terrestrial radiation (Lovis 1977; Schneider et al.
2004a; Rothwell & Stockey 2008; Schuettpelz & Pryer 2009; H.-M. Liu et al. 2014). Well
over 70% of ferns are eupolypods, and about one third of these are epiphytes, amd
these make up ca 10% of all vascular epiphytes. Diversification of epiphytic ferns in
particular occurred during the Palaeogene, perhaps linked with the Palaeocene-Eocene
thermal maximum (Schneider et al. 2004a, b; Schuettpelz 2007; esp. Schuettpelz &
Pryer 2009: Supplemental Tables 2, 3; Watkins et al. 2010; c.f. Dubuisson et al. 2009, in
part). An exception is Trichomanes and relatives (but not Hymenophyllum and its
relatives, for which, see Dubuisson et al. 2009) which diversified in the early Cretaceous;
they are commonly epiphytic on tree ferns, which themselves had begun diversifying in

the Jurassic (Schuettpelz 2007; also Schuettpelz & Pryer 2009; Rothwell & Stockey 2008
for early radiations of leptosporangiate ferns).
A provisional hierarchy of characters taken from Smith et al. (2006, 2008), Pryer et
al. (1996) and Rothfels et al. (2012b) is given below. For a first stab at apomorphies for
members of the polypod II clade, see Sundue and Rothfels (2013); their suggestions are
largely followed here; problems with sampling, character state delimitation, and different
apomorphies produced by different methods of character optimization are taken into
account as far as possible. Only unambiguous synapomorphies (Sundue & Rothfels 2013)
are flagged below. Kaplan (1997, vol. 3: chap. 19) summarized the sporangium wall
morphology of monilophyta; the annulus is an apomorphy of Polypodiopsida, the
leptosporangiate ferns. For other possible apomorphies, see e.g. Schneider et al. (2009).
More features need to be added, e.g. if the megaphyllous leaves of most ferns and those
of seed plants have evolved independently, which seems likely (see above).
Understanding the evolution of the apparently very unfern-like plant body of
Psilotales and of Psilotum itself, until fairly recently considered to be perhaps the most
primitive extant vascular plant - when I was taught about Psilotum, it was compared
with the Palaeozoic rhyniophytes - still presents difficulties. Kaplan (1997, vol. 3: chap.
10; Siegert 1973 and references) summarized the literature, noting that these problems
were because the morphology of Psilotum in particular was compared with these fossil
plants, now thought to be entirely unrelated. As he noted, young leaves of Psilotum did
have features of fronds (see also Kaplan 1977). Similarly, the leaves of Equisetum may
be secondarily simple. Thus its fossil relative Sphenophyllum has much larger leaves with
dichotomous venation; bifacial xylem has been found in this genus (Kenrick & Crane
1997; Doyle 2013 and references).
Ecology & Physiology. The evolutionary physiology of ferns is repaying examination.
The tracheids tend to be long and wide, and the scalariform perforation plates may
extend the length of the cell. As a result, water transport can be relatively efficient;
conifers, another group without vessels and that also has quite efficent water transport
at least has secondary thickening and bordered pits (Pittermann et al. 2011). Fern
richness in Australia is correlated with precipitation, and depending on what aspect of
diversity is emphasized, precipitation may be linked to equability, etc., bryophytes, not
not flowering plants, showing rather similar patterns (Nagalingum et al. 2015).
Epiphytic ferns are concentrated in Hymenophyllaceae, Polypodiaceae, Pteridaceae
(e.g. Vittaria) and Dryopteridaceae (e.g. Elaphoglossum) (Schuettpelz & Pryer 2009;
Kato & Tsutsumi 2013; see also above). The gametophytes of these epiphytes are often
strap-shaped and long-lived, and are notably resistant to dessication; as Watkins et al.
(2007: p. 716) observed, "fern gametophytes are, for all intents and purposes,
bryophytes", and the epiphytic species they studied were the most resistant (see also
Nayar & Kaur 1971: survey of gametophyte diversity; Dassler & Farrar 2001; Farrar et
al. 2008; Watkins & Cardels 2012; Rothfels & Schuettpelz 2014). Gemmae on these
gametophytes (e.g. Pryer et al. 2016) will further increase their longevity. Ferns show a
variety of other adaptations to the epiphytic habitat, including CAM photosynthesis
(Watkins & Cardels 2012) and production of gemmae (Farrar 1974), and mycorrhizae
are less frequent (e.g. B. Wang & Qiu 2006; Kato & Tsutsumi 2013); nutrients tend to be
low in the epiphytic habitat. Testo and Sundue (2014) found that both terrestrial species
and hemiepiphytes (where roots from the climbing plant establish a connection with the
soil) were derived within a clade of epiphytic Polypodiaceae, while J. Clark et al. (2016)
suggested that there may be a connection between the epiphytic habit and increased
genomes size in the eupolypods.
Kawai et al. (2003) found a distinctive chimaeric red/far red light photoreceptor (phy
3, = neochrome) in which red-sensing phytochrome and blue-sensing phototropin are

fused into a single molecule (F.-W. Li et al. 2014) in some Polypodiales, possibly aiding in
phototropic responses in the shaded conditions in which ferns have diversified;
neochrome is very sensitive to white light (Wada 2008). Neochrome seems to have been
acquired by polypod ferns from hornworts via lateral gene transfer around 179 m.y.a.,
which, perhaps not so coincidentally, is about the age of Polypodiales (Schneider et al.
2004a), and may even have been acquired more than once (F.-W. Li et al. 2014).
In seed plants, stomata open in response to both red and blue light. Stomata of
those few ferns examined, all leptosporangiates (Pteris, Adiantum, Asplenium and
Nephrolepis), lack a blue light-specific opening response, although elements of the
response, including the relevant phototropin, a blue right receptor protein kinase, are
present (Doi et al. 2006). In the shaded understory environments that many ferns
prefer, blue (and red) light has been preferentially absorbed by the canopy above;
stomatal closure is mediated by red light. Furthermore, ferns do not have leaves in which
the veins are the same distance from the epidermis as they are from each other; they
are not hydraulically optimised. However, photosynthesis may proceed best in the low
levels of diffuse light and so this hydraulic optimisation may not matter so much
(Zwieniecki & Boyce 2014). (In seed plants the effectiveness of the response is
white/blue + red > blue > red > green light [Willmer & Fricker 1996] and in many
flowering plants leaves are hydraulically optimised [Zwieniecki & Boyce 2014].)
Fertilization & Spore Dispersal. The distributions of the sporophytic and
gametophytic plants of the one fern species may be quite different, particularly if the
gametophyte is other than heart-shaped. Strap-shaped or filamentous gametophytes can
live for a very long time and/or produce gemmae, and so they can persist in sites that
are hundreds of miles from the nearest sporophytes (e.g. Farrar 1967; Ebihara et al.
2013). With the advent of the ability to identify gametophytes directly by molecular
sequencing, rather than waiting for them to produce sporophytes, this phenomenon is
turning out to be remarkably common (Ebihara et al. 2013), and it has interesting
implications for the evolution of ferns (see e.g. Ebihara et al. 2009).
Noblin et al. (2012) describe the mechanics of how the annulus functions in
sporangium dehiscence.
Plant-Animal Interactions. Overall, herbivory in ferns is about the same as the
average for angiosperms (Turcotte et al. 2014: see caveats).
Bacterial/Fungal Associations. For mycorrhizae in ferns, see Lehnert et al. (2010 and
references). Mycorrhizal associations are not known in Equisetum (Read et al. 2000).
Genes & Genomes. For the evolution of the monilophyte chloroplast genome, see
Karol et al. (2010), Grewe et al. (2013) and J. Clark et al. (2016). Within Polypodiopsida,
some inversions in the chloroplast inverted repeat may be high-level synapomorphies
(Gao et al. 2009). Grammitidaceae s. str. in particular (included in Polypodiaceae) have
green spores and accelerated plastid genome evolution, a correlation found elsewhere in
ferns, although it is not 100% (Schneider et al. 2004b), indeed, spores that less
obviously contain chloroplasts are quite widespread (Sundue et al. 2011).
Ferns are noted for their high incidence of polyploidy, almost 1/3 (31%) of all
speciation events being accompanied by polyploidy (Wood et al. 2009), and base
chromosome numbers are also usually very high. There has been recent hybridization
between clades (Cystopteris, Gymnocarpium) that diverged an estimated (76.2-)57.9(40.2) m.y.a. (Rothfels et al. 2015a) while there is hybridization in Osmunda between
clades that may have been separated for four times as long (Bomfleur et al. 2014b;
Grimm et al. 2015).

For information on genome size, see Obermayer et al. (2002) and J. Clark et al.
(2016); particularly large genomes occur in Psilotales and some Ophioglossales, and in a
few polypods.
Chemistry, Morphology, etc. The xyloglucan composition of the primary cell wall
varies substantially, that of Equisetum being particularly distinctive (Hsieh & Harris
2012). For cell wall polysaccharides, see also Silva et al. (2011).
Basic sporophyte morphology was outlined by Kaplan (1997, vol. 2: chap. 11, 18,
vol. 3: chap. 19, 2001). For the organization of the apical meristem, see Amrbose and
Vasco (2015). The stem has a siphonostele, the protoxylem being restricted to lobes of
the central xylem strand, hence bringing to mind a necklace (development of the xylem
is mesarch, although it is notably variable in the Ophioglossum/Psilotum clade). The
protoxylem is described as having G-type tracheids (Edwards 1993). HernndezHernndez et al. (2012) discuss the distribution of the circumendodermal band, tannincontaining cells surrounding the petiolar leaf trace that have a common orgin with the
endodermis; they also detail the distributions of a number of other vegetative/habit
features. Vasco et al. (2013) summarize fern leaf morphology and development, noting a
number of shoot-like features. Davies (1991) summarized information about aerophore
distribution in ferns. The information on horizontal cell walls in early embryo
development in ferns given by Philipson (1990) seems to be incorrect - the examples
should be vertical?
For the stem apex of ferns, see above.
Takahashi et al. (2009, 2014 and references) describe gametophyte development in
ferns. They note that the apical region converts to a multicellular meristem, which can
divide - dichotomous branching - if cell division in the middle of the meristem stops; the
branched, strap-shaped gametophytes of epiphytic ferns are simply an extreme variant
of this morphology. Archegonia develop only after the formation of the multicellular
meristem. For details of male gamete morphology and movement, etc., see e.g.
Renzaglia et al. (2000b, 2002) and Schneider et al. (2002).
For information on pteridophytes in general (these have often - and still may include lycophytes), see also Kato (2005) and Ranker and Haufler (2008). For other
general information, see Raven and Edwards (2001), for comparative anatomy, see
Ogura (1972), for vessels, see Sen and Mukhopadhyay (2014 and references), for
venation, see Wagner (1979) and Boyce (2005b), for details of stelar morphology and
evolution, see Beck et al. (1982), and for young sporophytes, etc., see Johnson and
Renzaglia (2009 and references).
Phylogeny. The circumscription of this clade has only recently become clear. It
includes the strongly supported [Psilotum + Ophioglossum] clade (Tmesipteris is sister
to Psilotum) perhaps sister to all other ferns, as chloroplast data has broadly tended to
suggest (Rothfels et al. 2015b for references). Wickett et al. (2014) obtained a
[Marattiales + Psilotales] clade sister to leptosporangiate ferns, but this may be a
sampling issue. The inclusion of morphology alone or in combination with molecular data
also affects the relationships detected (Wikstrm & Pryer 2005 and references); see also
Grand et al. (2013) for various morphological analyses. The position of Equisetum is
uncertain.
1. It may be sister to Angiopteris, etc. (although support only moderate), the combined
clade in turn being sister to remaining ferns (e.g. Pryer et el. 2001a, 2004a; Wikstrm &
Pryer 2005; Qiu et al. 2007; Ebihara et al. 2011; c.f. in part Wolf et al. 1998).

2. It may be sister to all other ferns, as in a rps4 analysis, and also 4- and 5-gene
analyses, the latter two with strong support (Schuettpelz et al. 2006), analyses of
several plastid genes (Rai & Graham 2010), and in a matK phylogeny (Kuo et al. 2011).
Knie et al. (2015) also find good support for the relationships [Equisetum [[Psilotum +
Ophioglossum] + The Rest]], as does Rothfels et al. (2015b) in their nuclear gene
analysis. Interestingly, spore wall ultrastructure of Calamites, an extinct equisetaceous
plant, is not so different from that of Ophioglossaceae and other ferns (Lugardon &
Brousmiche-Delcambre 1994; Grauvogel-Stamm & Lugardon 2009). Equisetum has no
mitochondrial atp1 intron, either a secondary (and parallel) loss or plesiomorphic
absence, depending on the topology of the whole group (Wikstrm and Pryer 2005: see
the character hierarchy below).
3. Schneider et al. (2009) noted potential morphological apomorphies such as simple
leaf blade and stems with both radial and dorsiventral symmetries (= erect plus creeping
stems...) suggesting a clade [Psilotales + Equisetales], consistent with some structural
changes in the chloroplast genome (Grewe et al. 2013; see also some analyses in Karol
et al. 2010; also Wolf & Karol 2012; Ruhfel et al. 2014; J.-M. Lu et al. 2015: chloroplast
genomes).
This reorganisation of monilophytes has sometimes been severely criticised
(Rothwell & Nixon 2006), but it is unclear how damning such criticism is. Since the
evaluation of "support" values for particular topologies is integral to the approach
adopted in these pages, the decision to exclude such values by those authors makes
their work difficult (for me, at least) to interpret. Indeed, in several morphological
cladistic analyses (e.g. Bremer 1985; Stevenson & Loconte 1996; Rothwell 1999: fossils
included or not) Psilotum came out as sister to all other vascular plants. However, some
morphological analyses (Schneider et al. 2009) do place Psilotum with other
monilophytes, even if the same analyses also place flowering plants within a paraphyletic
group of extant gymnosperms.
Classification. Smith et al. (2006, 2008) propose a phylogeny-based
reclassification of the ferns, and they also include literature, ordinal and familial
synonymy, and a list of accepted genera and some major synonyms; Prelli (2010) gives
a nice account of European ferns. However, adjustments to this classification are being
made as details of the phylogeny become better understood (Schuettpelz & Pryer 2007,
2008; Kuo et al. 2011; Rothfels et al. 2012b: reclassification of eupolypods II). A linear
sequence of families and genera (Christenhusz et al. 2011a) is now dated, but more
recently Christenhusz and Chase (2014) have proposed another classification. There are
some differences between the hierarchy below and that used by e.g. Rothfels et al.
(2015b), but they will be cleared up as things (hopefully) settle down.
Previous Relationships. Psilotum and Equisetum were previously thought to
represent independent lineages, with Psilotum and relatives considered to be the most
primitive living vascular plants, and the latter do look superficially similar to some early
fossils. Their association with ferns, now very largely accepted, was unexpected (but see
Kenrick & Crane 1997). Although Bierhorst (1968, see also 1977) compared Psilotum
with the extant fern Stromatopteris and found some morphological similarities, most of
these have turned out to be parallelisms and the two are not at all close.

EQUISETOPSIDA / [Equisetales [Psilotales + Ophioglossales]]: plant with erect and

creeping stems; ?vernation; tapetum plasmodial; embryo exoscopic, suspensor 0;
chloroplast rps16 gene and rps12i346 intron lost.
Age. The clade that contains Equisetum has probably been separate from other
monilophytes since the Permian, ca 250+ m.y.a. (Stanich et al. 2009); Zhong et al.
(2014b) thought that this clade was (370.3-)296.2(-189.9) m.y. old.

EQUISETALES Berchtold & Presl

EQUISETACEAE de Candolle

Plant with erect and creeping stems; roots triarch, with large central tracheid; cell wall
also with (1->3),(1->4)--D-MLGs [Mixed-Linkage Glucans], Si02 accumulation
common; stem with intercalary meristem [at base of leaf sheath], ridged,
photosynthetic, protoxylem mesarch, with central canal; protoxylem lacunae
developing; leaf vascular bundles amphicribral; branches whorled, members of whorls
alternating at each node; leaves small, simple, 1-veined, whorled, basally connate, not
photosynthetic; sporangiophores peltate, aggregated into a strobilus; sporangial cell
walls with helical secondary thickenings; tapetum plasmodial; spores with circular
aperture [hilate], abapertural obturator +, green, wall with silica, elaters 4-6/spore,
spatulate, helically-coiled; embryo exoscopic, plane of first cell division variable,
suspensor 0; n = 108; mitochondrial atp1 intron 0.
1/15. World-wide, not the Antipodes.
Age. Extant species of Equisetum seem to have separated in the Caenozoic
(77.5-)64.8(-52.1) m.y.a. (Des Marais et al. 2003, but c.f. Stanich et al. 2009).
Fossils with many of the apomorphies of crown group Equisetum are known from
Upper Jurassic deposits from Patagonia some 150 m.y. or more old (Channing et al.
2011; see also Stanich et al. 2009); for still older Equisetum-like spores - but with trilete
marks - and associated elaters from the Middle Triassic, see Schwendemann et al.
(2010).
Evolution. Divergence & Distribution. Change in spore morphology from the
Calamites type to the at first sight very different trilete spores of Equisetum is
convincingly demonstrated by Grauvogel-Stamm and Lugardon (2009). Other fossils
placed in this area include Sphenophyllum, which has megaphylls, and this suggests that
the tiny leaves of Equisetum may be reduced rather than being microphylls. Roots of
fossils also differ in morphology and can be polyarch.
Ecology & Physiology. Equistetum tends to grow in ecologically rather stressfull
habitats, including hot springs (Channing et al. 2011 and literature; Husby 2012). There
is pressurized gas flow, so-called convective ventilation, from the stems into the
rhizomes in some extant species of Equisetum and probably in their fossil calamitalean
relatives from the Carboniferous. Oxygen moves via interconnected air spaces into the
rhizomes, so allowing them to penetrate deeply into the anoxic substrates commonly
favoured by this group (Armstrong & Armstrong 2009). However, those species of
Equisetum that lack interconnected air spaces - and this is accompanied by changes in

position of the flow-resistant endodermis in the stem - have no convective ventilation

(Armstrong & Armstrong 2011).
Reproductive Biology. Aided by their elaters, spores of Equisetum can either jump up
to 1 cm in the air as they dry, or move by short random walk-type movements along the
ground (Marmottant et al. 2013).
Genes & Genomes. For a genome duplication in Equisetum dated to (112.5-)92.4(75.2) - or perhaps only 70-50 - m.y.a., see Vanneste et al. (2015).
Chemistry, Morphology, etc. For (1->3,1->4)--d-glucans, see Fry et al. (2008)
and Xue and Fry (2012), and for Si concentration, etc., see Husby (2012).
Phylogeny. Equisetum bogotense is sister to the rest of the genus (Guillon 2007).
[[Psilotales + Ophioglossales] [Marattiopsida + Polypodiopsida]]: (spore aperture
proximal and monolete); indel in mitochondrial rpl2 coding region.
Age. Rothfels et al. (2015b) suggested an age of (390.4-)351.7(-309.7) m.y. for
this node.
[Psilotales + Ophioglossales]: plant with erect and creeping stems; stem protoxylem
development variable; embryo exoscopic, suspensor 0; gametophyte subterranean,
axial, non-photosynthetic, mycorrhizal; nuclear genome size 32.7-64.8 pg.
Age. Magalln et al. (2013) estimated an age of around 275.6 m.y. for this clade,
and (316-)306(-267) m.y. is the age in Y. L. Qiu et al. (2007), ca 214.5 m.y. in Laenen
et al. (2014) and (317.1-)250.5(-141.8) m.y. in Rothfels et al. (2015b); see also P. Soltis
et al. 2002) for estimates.
Genes & Genomes. Both Psilotum and Ophioglossum have large genomes with
(Bennett & Leitch 2005; J. Clark et al. 2016).
PSILOTALES Prantl
PSILOTACEAE J. W. Griffith & Henfrey

Epiphytes; roots 0; ?leaf vascular bundles; leaves small, simple, (laterally flattened Tmesipteris), veins 1 or 0; sporangia 2-3, forming synangium; tapetum glandularamoeboid; spores kidney-shaped, monolete; gametophyte with septate rhizoids; (transfer
cells in sporophyte only - Psilotum), n = 52, 208, genome size [1C] = ca 72.5 pg. 2/12.
Age. Zhong et al. (2014b) estimated an age of (147.1-)72.3(-14.7) m.y. and
Rothfels et al. (2015b) an age of (142.5-)78.9(-28.5) m.y.a. for this clade.
OPHIOGLOSSSALES Link
OPHIOGLOSSACEAE Martinov

Root with 2-5 protoxylem poles; root hairs 0; cork mid cortical; vascular cambium +;
stem stele sympodial; tracheids with circular bordered pits; leaf vascular bundles
collateral; (axillary buds +); fronds compound to simple, 1 produced/year, venation
reticulate, with internally directed veins, vernation nodding, bases sheathing, stipules +,
thin; one or more sporophores associated with each tropophore; (gametophyte with
septate rhizoids); (embryo endoscopic; first cell wall of the zygote vertical); n = (44) 45
(46 ...720), genome size [1C] = ca 28.35 pg. 4/80.
Age. Rothfels et al. (2015b) suggested an age of (249.6-)161.7(-74) m.y. for
crown-group Ophioglossaceae.
Glomeromycote mycorrhizae in Ophioglossum are associated with the
echlorophyllous gametophyte and subterranean sporophytic stage, and also the
photosynthesising sporophyte (Field et al. 2015a). Takahashi and Kato (1988) describe
the development of lateral meristems in the family.
Ophioglossum reticulatum, at n = 720, has the highest chromosome numbers of any
plant.
See Hauk et al. (2003) for a phylogeny, Mankyua not included, also Shinohara et al.
(2013), Mankyua included, but position unstable - sister to rest of family (also in the
joint analysis), or to Ophioglossum s. str..
[Marattiopsida + Polypodiopsida]: leaf vascular bundles amphicribral; frond compound,
vernation circinate; scales +; sporangia abaxial; gametophyte green, surficial;
shoot developing towards the archegonial neck [from hypobasal cell, endoscopic];
nuclear genome 3.5-14.0 pg [?here]; changed gene adjacencies at borders of
chloroplast IR; mitochondrial atp1i361g2 intron gain.
Age. Zhong et al. (2014b) suggested an age of (378.8-)336.7(-291.5) and Rothfels
et al. (2015b) an age of (364.1-)329(-289.2) m.y. for this clade.
P. Soltis et al. (2002) offer a variety of suggestions for node ages within this clade.
MARATTIOPSIDA
Roots with several protoxylem poles; root hairs septate [?multicellular]; petiole
vasculature polycyclic; stipules +, fleshy and starchy.
MARATTIALES Link
Synonymy: Christenseniales Doweld
MARATTIACEAE Kaulfman

Dictyostele +; mucilage canals +; rhizome with scales; aerophores linear, with lenticels;
fronds pulvinate, (leaflets with internally directed reticulate venation - Christensenia);
meiosis monoplastidic [?all}; spores bilateral or ellipsoid, monolete; transfer cells 0; x =
40. 5/150: Danaea (50).
For meiosis, see Brown and Lemmon (2001).
Evolution. Divergence & Distribution. For some comments on biogeography, see
Christenhusz and Chase (2013). There seems to be a slow-down in the rate of evolution
in this clade (Rothfels et al. 2015b and references).
Phylogeny. For a phylogeny, see Murdock (2008a), also Christenhusz et al. (2008);
the fossil Psaronius seems to associate consistently with Marattia (e.g. Grand et al. 2013
and references).
Both Marattia and Angiopteris are paraphyletic, but they can easily be made
monophyletic (Murdock 2008b). For Danaea, see Christenhusz (2010).
POLYPODIOPSIDA / leptosporangiate ferns.
Primary cell walls poor in mannans and rich in tannins; roots with 2 protoxylem poles;
primary xylem with scalariform bordered pits; leaf trace single; aerophores linear, on
either side of the petiole, with stomata; sporangium derived from periclinal division of a
single epidermal cell, wall one-layered, stalk 4-6 cells across [= leptosporangiate];
sporangium with exothecium forming an annulus, 64-800 spores/sporangium;
antheridium exposed; gametophyte cordate [level?]; embryo prone [first cell wall of
the zygote vertical, parallel to gravity], with quadrant/octant formation, suspensor 0.
Age. Magalln et al. (2013: with temporal constraints) estimated an age of around
(267.8-)252.7-251.4(-246.1) m.y. for this clade; (330-)323(-310) m.y. is the age in Y. L.
Qiu et al. (2007), perhaps 350 m.y.a. in Schneider et al. (2004a), ca 299 m.y.a. in
Schuettpelz and Pryer (2009), and around 170 m.y.a. in Villarreal and Renner (2014)
and (327.8-)301.3(-271.5) m.y. in Rothfels et al. (2015b) - all told, a rather
disconcerting spread of ages.
Phylogeny. The large clade made up of leptosporangiate ferns has very strong
support (see also Hasebe et al. 1994, 1995, Pryer et al. 1995; Wolf et al. 1998; Quandt
et al. 2004; Schuettpelz et al. 2006; Rai & Graham 2010, etc.). Rothfels et al. (2015b)
emphasized that their analyses of nuclear data broadly agreed with several plastid
sequence analyses.
Within the leptosporangiates, Osmunda and relatives, the sporangia of which have
some eusporangiate features, are strongly supported as being sister to the rest. There is
further substantial resolution of relationships (e.g. Pryer et al. 2004a, b and references;
Schuettpelz & Pryer 2007; c.f. in part Kuo et al. 2011: positions of Gleicheniaceae,
Lindsaeaceae, Nephrolepis [previously in Lomariopsidaceae] uncertain). It is unclear
whether or not there is a clade [Hymenophyllales + Gleicheniales] (Knie et al. 2015 for
literature; Rothfels et al. 2015b: Gleicheniales paraphyletic, but with little support). J.-M.
Lu et al. (2015: chloroplast genome, but sampling) found Dipteridaceae and Lygodiaceae
to be successive branches along the leptosporangiate stem. Schizaeales and Salviniales
(strong support) and Cyatheales (weak support) are successively sister to Polypodiales

(e.g. Rothfels et al. 2015), and Rai and Graham (2010) suggest additional variants. For
relationships with Schizaeales, see Labiak et al. (2015 and references).
Within Polypodiales, Lindsaeaceae are probably sister to all others (see also Rothfels
et al. 2015b), but the genera Cystodium, ex Dicksoniaceae, and Lonchitis and
Saccoloma, both ex Dennstaediaceae - the last as a separate family below - are also in
this area (Lehtonen et al. 2012). Pteridaceae and Dennstaediaceae were well supported
as successive sister taxa to the eupolypods (Rothfels et al. 2015b; J.-M. Lu et al. 2015).
Davalliaceae and related taxa are sister to the polygrammoid ferns, and both include a
number of epiphytes (for their evolution, see Tsutsumi & Kato 2006). Relationships
suggested by structural changes in the chloroplast genome (Wolf & Roper 2008; Wolf et
al. 2010, 2011) are consistent with those suggested by sequence analyses. Cystopteris
and relatives form a clade that may be sister to the eupolypod II clade (Rothfels et al.
2009, esp. 2012a, 2013, 2015b).
OSMUNDALES Link
OSMUNDACEAE Martinov

Cataphylls [petiole bases] +; Si02 accumulation common; stem with ectophloic

siphonostele, with a ring of conduplicate/twice conduplicate discrete bundles; leaves with
stipules; fronds with fertile and sterile portions (fertile and sterile fronds separate);
annulus a lateral group of cells; spores green; (rhizoids septate), zygote elongating; n =
22. 4/20.
The Osmunda clade originated in the late Carboniferous, ca 323 or 305 m.y.a.
(Phipps et al. 1998; Schneider et al. 2004a), and is very diverse from the Permian
onwards, less so more recently. Fossils some 180 m.y.o. have anatomy that is
remarkably like that of the extant Osmunda claytoniana, and probably the chromosome
number and genome size of the fossil plant were similar, too (Bomfleur et al. 2014a).
Schneider et al. (2015) reconstruct the history of genome size changes in Osmundaceae;
there is some variation. Generation times in royal ferns are long, so genome change
might be expected to be low, but whether or not there is "genomic stasis" (Bomfleur et
al. 2014a; c.f. Schneider et al. 2015) is another issue; there has been some change.
Age. The age of this clade is estimated at around 199.6 m.y. (Schuettpelz & Pryer
2009); however, estimates from Carvalho et al. (2013) based on fossils that can be
assigned to the leptosporangioid branch of the tree, suggest an age in excess of ca 265
m.y.a., Late Permian (see also Wilf & Escapa 2014), while the preferred age in Grimm et
al. (2015: comprehensive analysis, also incorporating fossils) is (264-)243(-233)
m.y..ago.
It has been suggested that Osmunda is paraphyletic, with Osmunda (now =
Osmundastrum) cinnamomea being sister to the rest of the family (Metzgar et al. 2008);
relationships in Carvalho et al. (2013) are [Osmundastrum [Osmunda [Leptopteris,
Todea, Todites]]]. However, Bomfleur et al. (2014b) argue for the monophyly of
[Osmunda + Osmundastrum] based on extensive data from fossils and a re-evaluation
of the molecular evidence, i.a. they note that O. cinnamomea is able to hybridize with
some species of Osmunda, but not with other Osmundaceae. Given an estimated date of
the split of the first two of (264-)238(-233) m.y.a., only a little less than the age of

crown-group Osmundaceae as a whole (Grimm et al. 2015), they have been separated
for far longer than any other vascular plants that hybridize.
[[Hymenophyllales + Gleicheniales] [Schizaeales [Salviniales [Cyatheales +
Polypodiales]]]]: protostele +; sporangia in sori, annulus oblique, continuous; loss of
chloroplast trnK gene and its intron.
Age. (297-)286(-272) m.y. is the age for this node in Y. L. Qiu et al. (2007), ca
280.1 m.y. in Schuettpelz and Pryer (2009: Hymenophyllales sister to the rest)
(306.8-)278.7(-252.3) m.y. in Rothfels et al. (2015b).
[Hymenophyllales + Gleicheniales]: ?
Age. The age for this node in Y. L. Qiu et al. (2007) was estimated at (283-)273(259) m.y. and in Rothfels et al. (2015b) at (276.7-)237.2(-192.4) m. years.
HYMENOPHYLLALES A. B. Frank
HYMENOPHYLLACEAE Martinov

Epiphytes common; axillary buds +; fronds 1 cell thick between veins, stomata 0;
sporangia on elongated receptacle, maturation basipetal; spores globose, green;
gametophyte filamentous or ribbon-like; embryo not with tetrad/octant formation; x =
36. 9/600.
Age. Crown Hymenophyllaceae are (190.4-)185.1(-174.7) m.y.o. (Schuettpelz &
Pryer 2009).
For the rate of molecular evolution if Hymenophyllaceae, with an apparent slowdown in Hymenophyllum, see Schuettpelz and Pryer (2007).
Around half the family is epiphytic (Zotz 2013), and there are also climbing taxa
(see Dubuisson et al. 2009 for growth forms in Hymenophyllum). Epiphytism in
Trichomanes evolved before that in Hymenophyllum, the plants probably growing on the
stems of Cyantheaceae on which species are still often to be found (Hennequin et al.
2008), indeed, diversification in Trichomanes is estimated to have begun in the middle of
the Jurassic and that in Hymenophyllum in the middle of the Cretaceous (Schuettpelz &
Pryer 2007), ca 147.3 versus ca 41.9 m.y.a. (Schuettpelz & Pryer 2009). See also
Schuettpelz and Pryer (2007) and Hennequin et al. (2008) for other dates.
Despite the delicate fronds of Hymenophyllaceae, dessication tolerance is at least
sometimes well developed - rather like mosses (Proctor 2003, 2012). Indeed, the
sporophytes of some epiphytic trichomanoid ferns have lost both cuticle and roots
("regressive evolution" - Dubuisson et al. 2011), and may be functionally similar to
bryophytes; the stem stele may have just a single vascular element (Dubuisson et al.
2013; see also Dubuisson et al. 2003b).
For the phylogeny of the family, see Pryer et al. (2001b) and Dubuisson et al.
(2003a, 2013), for that of Trichomanes and relatives, see Ebihara et al. (2007), for that
of Hymenophyllum, with a focus on the large subgenus Mecodium, which turns out to be

polyphyletic but common in a number of basal clades, see Hennequin et al. (2006), and
for a possible base chromosome number in the family - previous suggestions x = 6-9,
11, 13, but here = 36 - see Hennequin et al. (2010).
[Gleicheniales + The Rest]: (spores monolete, perine closely attached ro exine).
Age. The stem group age of this node is ca 276.4 m.y. (Schuettpelz & Pryer 2009).
GLEICHENIALES Schimper
Root steles with 3-5 protoxylem poles; rhizome with scales; frond veins anastomosing;
sporangium maturation simultaneous; (gametophyte axial - Stromatopters), (rhizoids
septate); antheridia with 6-12 narrow curved or twisted cells in walls; x = 20, ... 116,
genome size [1C] = 2.96 pg [sampling].
Age. Crown-group Gleicheniales are ca 262.2 m.y. (Schuettpelz & Pryer 2009) or
(252.4-)196.1(-134) m.y.o. (Rothfels et al. 2015b).
Synonymy: Dipteridales Doweld, Matoniales Reveal, Stromatoperidales Reveal
GLEICHENIACEAE C. Presl

Leaves indeterminate, pseudodichotomously forked (not - Stromatopteris); spores

(bilateral), monoulcerate; (embryo exoscopic, cell wall vertical, gametophyte (axial,
subterranean), with clavate hairs; x = 22, 34, etc.. 6/125.
There has been a chloroplast genome inversion in the family (Wolf & Roper 2008).

[Dipteridaceae + Matoniaceae]: ?
DIPTERIDACEAE Seward & E. Dale

Frond veins reticulate, areoles with included veins, veins 4.4-5.6 mm/mm2; sporangia
with "long" stalks, (spores bilateral, monolete); x = 33. 2/11. N.E. India to N.E. Australia,
earlier in Caenozoic widespread.

MATONIACEAE C. Presl

Stems solenostelic, with two vascular cylinders and a central bundle; fronds or pinnae
dichotomously branched; sporangia in ring surrounding central "receptacle", sorus
indusiate; x = 25, 26. 2/4. Malesia, previously widespread.

[Schizaeales [Salviniales [Cyatheales + Polypodiales]]]: plant with hairs; endospore 2layered; antheridium wall ca 3 cells across; two overlapping inversions in chloroplast
genome.
Age. (281-)266(-250) m.y. is the age for this node in Y. L. Qiu et al. (2007), ca
264.6 m.y. in Schuettpelz and Pryer (2009) and (289.4-)258.3(-235.2) m.y. in Rothfels
et al. (2015b).
SCHIZAEALES Schimper
Fronds differentiated into fertile/sterile portions [hemidimorphic]; annulus sub-apical; n
= 28-504.
Age. The crown-group age of Schizaeales is estimated to be around 218.4 m.y.
(Schuettpelz & Pryer 2009).

LYGODIACEAE M. Roemer

Fronds indeterminate, climbing, pseudodichotomously branched, with a bud in

angle of branch; one sporangium/sorus, subtended by antrorse indusium-like flange; x
= 29, 30. 1/25.

[Anemiaceae + Schizaeaceae]: sporangia not in sori, exindusiate.

ANEMIACEAE Link

Rhizome with dictyostele or solenostele, (with pockets axillary to the fronds); spores
tetrahedral, with parallel solid ridges (ridges hollow, centre spongy); x = 38. 1/100.
For a largely well-resolved phylogeny of the family and optimisation of characters on
to the tree, see Labiak et al. (2015: spore morphology!).

SCHIZAEACEAE Kaulfman

Inner pericyclic cells 6, 8, thickened; fronds undivided or fan-shaped, veins

dichotomizing; sporangia borne on marginal projections at blade tip; spores
monolete, bilateral; gametophyte filamentous, (white, subterranean, tuberous), rhizoids
septate, (embryo exoscopic, cell wall vertical); x = 77, 94, 103. 2/30.
There has been a chloroplast genome inversion somewhere around here (Wolf &
Roper 2008); see also the next node up.

[Salviniales [Cyatheales + Polypodiales]]: sporangium stalk 1-3 cells across [?position];

two [more!] overlapping inversions in chloroplast genome.
Age. The age of this node is estimated to be around 234.7 m.y. (Schuettpelz &
Pryer 2009) and (269.1-)231.6(-190.8) m.y. (Rothfels et al. 2015b).
Genes & Genomes. The rate of evolution of the 18S nuclear gene was lower than in
the other vascular plants examined (Stenien 2008: lycophytes not included).
SALVINIALES Bartling
Aquatics, aerenchyma +; stems dichotomizing; veins anastomosing;
sterile/fertile frond dimorphism; plant heterosporous, sporangia lacking
annulus; megaspore single; gametophyte development endosporic; genome size
[1C] = ca 2.38 pg; nrDNA with 5.8S and 5S rDNA in separate clusters.
Age. Crown-group Salviniales are estimated to be ca 186.8 m.y.o. (Schuettpelz &
Pryer 2009).
See Nagalingum et al. (2006, 2007: sporocarp structure) and Nagalingum et al.
(2008: phylogeny).
Synonymy: Marsileales von Martius, Pilulariales Berchtold & Presl
MARSILEACEAE Mirbel

Leaves simple, or to 4 leaflets/frond; sori in stalked bean-shaped sporocarps [folded

pinnae]; megaspore with acrolamella over the exine aperture, perine gelatinous;
female gametophyte with 1 archegonium; n = 10, genome size [1C] ca 0.8 pg [Azolla].
3/75.
For the phylogeny of Marsilea and character evolution there, see Nagalingum et al.
(2007).
SALVINIACEAE Martinov

Plant free-floating; fronds sessile, 2-ranked, <2.5 cm long, simple; n = 9, 22. 2/16.
Azolla has Nostoc in its tissues and is an important nitrogen fixer in rice paddies,
etc..
[Cyatheales + Polypodiales]: dictyostele +; hydathodes +; IR with several large
inversions, ycf2 duplication.
Age. The age for this node is ca 211 m.y. in Y. L. Qiu et al. (2007), ca 223.2 m.y. in
Schuettpelz and Pryer (2009), (270.1-)228.8(-187.5) m.y. in Zhong et al. (2014b) and
(238.1-)204.6(-179) m.y. in Rothfels et al. (2015b).

CYATHEALES A. B. Frank
Hairs +; sori terminal on veins, indusiate, indusium with outer and inner parts;
sporangium stalk ca 5 cells across; antheridium walls 5 cells across; genome size [1C]
= ca 7.91 pg [?sampling].
Age. The crown-group age of Cyatheales is ca 186.7 m.y. (Schuettpelz & Pryer
2009) or rather younger, (167.8-)109.1(-56.8) m.y. (Rothfels et al. 2015b: Alsophila
sister to the rest).
There seems to be a slow-down in the rate of evolution at the base of Cyatheales
(Rothfels et al. 2015b and references).
Synonymy: Dicksoniales Reveal, Hymenophyllopsidales Reveal, Loxsomatales
Reveal, Metaxyales Doweld, Plagiogyriales Reveal
THYRSOPTERIDACEAE C. Presl

Indusium cup-shaped, receptacle columnar, clavate; x = ca 78. 1/1.

[[Loxsomataceae [Culcitaceae + Plagiogyriaceae]] [Cibotiaceae + Cyatheaceae +

Dicksoniaceae + Metaxyaceae]]: ?
[Loxsomataceae [Culcitaceae + Plagiogyriaceae]]: ?
LOXSOMATACEAE C. Presl

Indusium urceolate, receptacle elongate, often exserted; gametophyte with scale-like

hairs; x = 46, 50. 2/2.

[Culcitaceae + Plagiogyriaceae]: ?
CULCITACEAE Pichi Sermolli

Petiole with omega-shaped [] bundle, open end adaxial; outer indusium scarcely
differentiated; sori with paraphyses; x = 66. 1/2.

PLAGIOGYRIACEAE Bower

Multiple leaf traces coming from a U-shaped bundle; young fronds with dense,
pluricellular, mucilage-secreting hairs; indusium 0; x = ?66. 1/15.

[Cibotiaceae + Cyatheaceae + Dicksoniaceae + Metaxyaceae]: paraphyses +.

CIBOTIACEAE Korall

Multiple leaf traces coming from a U-shaped bundle; stomata with three subsidiary cells;
spores with equatorial flange, usu. parallel ridges on distal face; x = 68. 1/11.
[Cyatheaceae + Dicksoniaceae]: the crown-group age of this clade, if it exists, is ca 150
m.y.o. (Janssen et al. 2008).
CYATHEACEAE Kaulfman

Stem with polycyclic dictyostele; multiple leaf traces coming from a U-shaped bundle;
scales large (also small); fronds large; indusium 0 to completely surrounding sporangia);
x = 69. 5/600.
Age. Crown-group Cyatheaceae are estimated to be ca 96 m.y.o. (Janssen et al.
2008).
The rate of chloroplast genome evolution has slowed down considerably here,
probably because of the long generation time of tree ferns (P. Soltis et al. 2002; esp.
Zhong et al. 2014b).
Korall and Pryer (2014) outline major biogeographic patterns in the group; initially
Gondwanan vicariance seems to be involved, although crown-goup diversification did not
begin until ca 96 m.y.a. with subsequent rather limited (for ferns) transoceanic long
distance dispersal. The 100+ endemic species of Cyathea on Madagascar may represent
a Pliocene ([4.24-]3.07> m.y) diversification of three separate clades each of which has
a fairly lengthy sojourn on the island - thus there was around 30 m.y. in one Malagasy
clade before diversification (Janssen et al. 2008, see also Korall & Pryer 2014).
Bystriakova et al. (2011) discussed niche evolution.
See Korall et al. (2006, 2007) for a phylogeny.
DICKSONIACEAE M. R. Schomburgk

Adaxial [outer!?] valve of sorus formed by reflexed frond segment margin and often
differently coloured from the other; x = 56, 65. 3/30.

METAXYACEAE Pichi Sermolli

Indusium 0; x = 95, 96. 1/2.

POLYPODIALES Link

Rhizome dorsiventral [?level]; sporangial maturation mixed; stalk 1-3 cells thick, annulus
vertical, interrupted by stalk and stomium; neochrome/phy 3 +.
Age. This clade is estimated to be some (200-)176(-163) m.y.o. (Schneider et al.
2004a), ca 191 m.y.o. (Schuettpelz & Pryer 2009) or (220.1-)184.2(-149.2) m.y.
(Rothfels et al. 2015b).
Synonymy: Aspleniales Reveal, Athyriales Schmakov, Blechnales Reveal,
Dennstaedtiales Doweld, Dryopteridales Schmakov, Lindseales Doweld,
Negripteridales Reveal, Parkeriales A. B. Frank, Platyzomatales Reveal, Pteridales
Doweld, Thelypteridales Doweld
LINDSAEACEAE M. R. Schomburgk

Innermost cortical layer of root usu. of 6 large cells; stele protostelic, with internal
phloem; leaf traces two, from V-shaped bundle; indusium opening towards margin; x =
34, 38, etc. 6/200. Pantropical (subtropical).
See Lehtonen et al. (2010) for a phylogeny and generic classification.
Age. (193.6-)165.4(-113.7) m.y. is suggested to be the age of this node in Rothfels
et al. (2015b).
CYSTODIACEAE J. R. Croft

1/.
LONCHITIDACEAE M. R. Schomburgk

1/.
SACCOLOMATACEAE Doweld

Scales?; petiole with omega-shaped [] bundle, open end adaxial; spores also with
distinctive parallel branched ridges; x = ca 63. 1/12.
[Dennstaedtiaceae + Pteridaceae]: Si02 accumulation common.
Age. Zhong et al. (2014b) suggested an age of (217-)154.3(-93.1) m.y. for this
node ([Denn. + Pterid.]), Schuettpelz and Pryer (2009) an age of ca 110.8 m.y., and
Schneider at al. (2016) ages somewhere around 137.4-95.6 m. years.

DENNSTAEDTIACEAE Lotsy

Chimaeric red/far red light photoreceptor [phy 3, neochrome]; stele?; hairs jointed;
petiole bearing buds, with omega-shaped [] bundle, open end adaxial;; x = 26, 29.
11/170.
Age. Crown-group Dennstaedtiaceae are estimated to be ca 72.2 m.y.o.
(Schuettpelz & Pryer 2009) or around 98-66.2 m.y.o. (Schneider et al. 2016).

PTERIDACEAE E. D. M. Kirchn.

(Epiphytic), (xeric); (petiole with omega-shaped [] bundle, open end adaxial;) indusium
0; (spores bilateral); (gametophyte ribbon-like); x = 29, 30. 50/950: Pteris (200-250),
Adiantum (200), Vittaria (80).
Age. This clade is around 106.3 m.y.o. (Schuettpelz & Pryer 2009), 90 m.y.o.
(Rothfels & Schuettpelz 2014), or about 99.9-87.4 m.y.o. (Schneider et al. 2016).
Kramopteris resinatus, in amber ca 100 m.y.o. from Myanmar, is placed at the split
between Monachosorum and other Pteridaceae (Schneider et al. 2016).
Phylogeny. For phylogenies, see Crane et al. (1995), Prado et al. (2007),
Schuettpelz (2007), Chao et al. (2014: Pteris, position of P. longifolia, the type, unclear)
and L. Zhang et al. (2015: Pteris somewhat expanded). Pryer et al. (2016) found that
Adiantum is monophyletic and is sister to Vittaria and its relatives. For increased rates of
molecular evolution, see Rothfels and Schuettpelz (2014). Cheilanthoid ferns, some 400
or more species, can grow in very dry conditions; generic limits are difficult here, but see
also Grusz et al. (2014 and references) and Yesilyurt et al. (2015).
Eupolypods: spores monolete, reniform, perine distinct; x = 41.
Age. This clade, which includes most ferns, is (124-)105(-91) m.y.o. (Schneider et
al. 2004a) or ca 116.7 m.y.o. (Schuettpelz & Pryer 2009).
[Didymochlaenaceae [Hypodematiaceae [[Lomariopsidaceae + Nephrolepidaceae]
[Dryopteridaceae [Tectariaceae [Oleandraceae [Davalliaceae + Polypodiaceae]]]]]]] /
eupolypod I: rhizome scales persistent, dense; leaf traces several, from V-shaped
bundle; petiole bundles several, circular [in t.s.], the two adaxial ones enlarged;
perispore with thick tuberculate folds/wings.
Age. The age of this node is ca 98.9 m.y. (Schuettpelz & Pryer 2009).
This is a largely epiphytic clade (Schuettpelz & Pryer 2009). For rhizome scales,
perhaps protecting the plant against dessication and aiding in the absorbtion of water
and nutrients, see Tsutsumi and Kato (2008); if they are not at this position on the tree,
they should be placed at the next node up (with parallel evolution within
Dryopteridaceae).

For relationships, see L.-B. Zhang and Zhang (2015); a couple more families may
still be needed. Thus H.-M. Liu et al. (2014), as they placed the odd genera Pteridrys and
Pleocnemia in Tectariaceae and Dryopteridaceae respectively, recognised an
[Arthropteridaceae + Tectariaceae] clade (the former is synonymized under the latter
here), but relationships are as suggested above.
DIDYMOCHLAENACEAE L.-B. Zhang & L. Zhang

Rhizome erect, subarborescent; sori hippocrepiform, somewhat elongated. 1/?1.

Pantropical.
[Hypodematiaceae [[Lomariopsidaceae + Nephrolepidaceae] [Dryopteridaceae
[Tectariaceae [Oleandraceae [Davalliaceae + Polypodiaceae]]]]]]: ?
HYPODEMATIACEAE Ching

(n = 40).
[Lomariopsidaceae + Nephrolepidaceae]: ?
LOMARIOPSIDACEAE Alston

?
NEPHROLEPIDACEAE

?
[Dryopteridaceae [Tectariaceae [Oleandraceae [Davalliaceae + Polypodiaceae]]]]: ?
DRYOPTERIDACEAE Herter

(Epiphytic); perine winged; gametophyte strap-like; x= 41. 30-35/1700: Elaphoglossum

(600), Polystichum (500). World-wide.
Age. Crown-group Dryopteridaceae are around 81.8 m.y.o. (Schuettpelz & Pryer
2009) or (94-)76(-58) m.y.o. (Le Pchon et al. 2016).
Elaphoglossum is the major epiphytic genus in the family - ca 400 species (ca 3/4 of
the genus) are epiphytes (Zotz 2013). Diversification within the speciose Polystichum
began in the Eocene, (60-)46(-32) m.y.a. (Le Pchon et al. 2016), that in
Elaphoglussum began a bit more recently - that covers about 2/3 of the family.
For a phylogeny of the whole family, see H.-M. Liu et al. (2007, esp. 2015); three
subfamilies are recognised, although two genera are unplaced. For a phylogeny of
Elaphoglossum, see also Lriga et al. (2014: 20-15 m.y.a. fossil of ?crown-group

Elaphoglossum from Dominican amber). Rouhan et al. et al. (2004) and Vasco et al.
(2015) discuss relationships within Elaphoglossum, where the Coast Rican E.
amygdalifolia and the Cuban E. wrightii are successively sister to the rest of the genus.
Moran et al. (2010a, b) investigate relationships within the bolbitidoid ferns focussing on
variation in perispore morphology. Li and Lu (2006a, b), L.-B. Zhang et al. (2012), Sessa
et al. (2012a), and McHenry and Barington (2014: exindusiate Andean species
monphyletic, sister to Mexican spp.) have been working on relationships within
Dryopteris itself, a genus whose limits are being clarified (e.g. Zhang & Zhang 2012) and
which shows extensive hybridization at all levels (Sessa et al. 2012b). Le Pchon et al.
(2016) examined phylogenetic and biogeographic relationships in Polystichum, while
Labiak et al. (2014) looked at relationships around Lastreopsis, with movement to and
fro between Australia and South America towards the middle of the Caenozoic.
[Tectariaceae [Oleandraceae [Davalliaceae + Polypodiaceae]]]: frond veins free, parallel
or pinnate.
TECTARIACEAE Panigrahi

(Climbers); (rhizome slender), (stipe and pinnae articulated); (frond veins free, parallel or
pinnate), with jointed usually stubby hairs; n = 40-42: Tectaria (200). 8-19/320.
Pantropical, inc. oceanic islands.
Age. Crown-group Tectariaceae (excluding Arthropteris) are around 50.4 m.y.o.
(Schuettpelz & Pryer 2009).
Although in the analysis of H.-M. Liu et al. (2013) the position of Arthropteridaceae
was uncertain, F. G. Wang et al. (2014a) found that they were embedded in
Polypodiaceae-Tectarioideae, as were Lomariopsidaceae; the latter was in a separate
clade in Schuettpelz and Pryer (2009). L.-B. Zhang and Zhang (2014) placed
Arthropteridaceae sister to Tectariaceae, but with less than overwhelming support,
Lomariopsidaceae were again separate.
Synonymy: Arthopteridaceae H. M. Liu, Hovenkamp & H. Schneider
[Oleandraceae [Davalliaceae + Polypodiaceae]]: fronds abscising from rhizome.
OLEANDRACEAE Pichi Sermolli

Fronds abscising just above the base [so leaving phyllopodia]; x = 41. 1/40.

[Davalliaceae + Polypodiaceae]: epiphytes predominant; perispore thin, verrucate,

granulate, etc..
The two diverged ca 42 m.y.a. (Sundue et al. 2015).
DAVALLIACEAE M. R. Schomburgk

Spores warty, warts close, not constricted at their bases [= verrucate-colliculate]; x = 40.
4-5/65.
Age. Crown-group Davalliaceae are ca 19.4 m.y.o. (Schuettpelz & Pryer 2009).
F.-G. Wang et al. (2014b) optimize spore morphology on a phylogeny of the family.
For a generic classification, see Kato and Tsutsumi (2008).
POLYPODIACEAE J. Presl & C. Presl

(Rhizome polysmmetrical); (Petiole with one or two vascular bundles - grammitids);

indusium 0; (spores green, globose-tetrahedral, trilete - grammitids); (gametophyte straplike); x = 35-37. 56/1200: Drynaria (50).
Age. Crown-group Dryopteridaceae (Loxogramme + The Rest) are about 55.8
m.y.o. (Schuettpelz & Pryer 2009).
For ages of splits of clades, see Schuettpelz and Pryer (2009) and Sundue et al.
(2015). Ca 87% of the species of Polypodiaceae are epiphytic (Zotz 2013), making them
the major epiphytic clade in the monilophytes, and diversification is associated with the
uplift of the Andes (e.g.), clades with broad elevational ranges diversifying faster,
speciation being related to climatic/environmental factors, less obviously associated with
morphological features (Sundue et al. 2015: epiphytes in general; Kreier et al. 2008:
Andean Serpocaulon; L. Wang et al. 2012: Qinghai-Tibet Lepisorus). Weber and Agrawal
(2014) suggested that the evolution of extra-floral nectaries in Pleopeltis was associated
with an increase in diversification.
For a phylogeny of microsoroid ferns, see Kreier et al. (2008), for that of grammitid
ferns, see Sundue et al. (2010, 2015 and references: generic changes). For the
phylogeny of grammitid ferns, see Sundue et al. (2014a, esp. b: Grammatis still
polyphyletic).
Janssen et al. (2005) discussed the evolution of the diverse frond morphologies in
Drynaria s.l.. For root anatomy, see Schneider (1996, 1997), and for petiole anatomy,
see Sundue et al. (2014a, esp. b and references).
F.-G. Wang et al. (2014a) include a broadened but monophyletic Tectariaceae as a
subfamily of Polypodiaceae.
[Cystopteridaceae [[Rhachidosoraceae [Diplaziopsidaceae [Desmophlebiaceae
[Hemidictyaceae + Aspleniaceae]]]] [Thelypteridaceae [Woodsiaceae [Athyriaceae
[Blechnaceae + Onocleaceae]]]]]] / eupolypod II: leaf traces two, from V-shaped
bundle, circumendodermal band surrounding trace; petiole bundles two,
elongated/crescent-shaped [in t.s.]; rhachis sulcus wall confluent with the costa of
pinna.
Age. This node is ca 103.1 m.y.o. (Schuettpelz & Pryer 2009).
CYSTOPTERIDACEAE Schmakov

Rhizome long-creeping; veins reaching the frond margin; indusium 0 or hood-like; n =

40, etc.. 3/36: Cystopteris (27).
For phylogenetic relationships, see Rothfels et al. (2013). The genera are
monophyletic, but there has been very extensive hybridization within Cystopteris and
Gymnocarpidium (Rothfels et al. 2014).
[[Rhachidosoraceae [Diplaziopsidaceae [Desmophlebiaceae [Hemidictyaceae +
Aspleniaceae]]]] [Thelypteridaceae [Woodsiaceae [Athyriaceae [Blechnaceae +
Onocleaceae]]]]]: ?
[Rhachidosoraceae [Diplaziopsidaceae [Desmophlebiaceae [Hemidictyaceae +
Aspleniaceae]]]] : sorus on one side of the vein.
RHACHIDOSORACEAE X. C. Zhang

Rhizome scales clathrate; petiole bundels U-shaped; n = 41; 1/4-7.

[Diplaziopsidaceae [Desmophlebiaceae [Hemidictyaceae + Aspleniaceae]]]: rhizome

scales not clathrate; scales on mature fronds 0.
DIPLAZIOPSIDACEAE X. C. Zhang & Christenhusz

Roots pale, fleshy, proliferous [producing plantlets by asexual reproduction]; fronds soft
and fleshy; vein endings raised and thickened, forming a submarginal vein; sori
elongated, only on one side of vein; n = 40, 41; 3/5.
[Desmophlebiaceae [Hemidictyaceae + Aspleniaceae]]: ?
DESMOPHLEBIACEAE Mynssen, A. Vasco, Sylvestre, Moran & Rouhan

Rhizomes erect or decumbent; petiole bundles hippocampiform, U-shaped or free; fronds

unequal-pinnate, veins free, forming submarginal vein; sori elongated; spores cristate; n =
? 1/2. Costa Rica to Brazil.
For information on Desmophlebiaceae, see Mynssen et al. (2016).
[Hemidictyaceae + Aspleniaceae]: ?
HEMIDICTYACEAE Christenhusz & H. Schneider

Fronds unequal pinnate, with submarginal collecting vein and margins with broad
membranous border; vein endings raised and thickened; n = 31. 1/1: Hemidictyum
marginatum. S. Mexico to S.E. Brazil.

ASPLENIACEAE Newman

Epiphytes common; root pericyclic sclereids with excentric lumina; leaf trace single,
circumendodermal band surrounding trace 0; petiole bundles back-to-back, C-shaped,
fusing and becoming X-shaped (V- or U-shaped); rhizome scales clathrate; frond usu.
with small clavate hairs; leaf margins decurrent and forming the lateral ridge along
rhachis; sori linear; indusia lateral, linear; sporangium stalk 1 cell across in the middle;
spores with decidedly winged perine; x = 35, 36, 38, 39. 1-10/700. Widely distributed.
Asplenium s.l. includes a large number of epiphytic species (Zotz 2013). Helical,
non-lignified wall thickenings (c.f. the velamen of monocots) occur in cortical cells of
some Asplenium, mostly epiphytic species (Leroux et al. 2011).
Age. Crown-group Aspleniaceae are ca 57.7 m.y.o. (Schuettpelz & Pryer 2009).
For generic limits, see Bellefroid et al. (2010 and references); Asplenium s. str. is
paraphyletic.
[Thelypteridaceae [Woodsiaceae [Athyriaceae [Blechnaceae + Onocleaceae]]]]: (Si02
accumulation common [?Athy., Onocl.]); frond once-pinnate/pinnatifid.
THELYPTERIDACEAE Pichi-Sermolli

Petiole vascular bundles uniting distally into a gutter shape; leaf hairs acicular, whitish or
hyaline, also on surface and/or margins of rhizome scales; first venation of frond
lobe/pinnule develops on basiscopic side [catadromous]; n = 27-36. 8/950: Microsorium
(600).
Age. The age of crown-group Thelypteridaceae is ca 68.5 m.y. (Schuettpelz & Pryer
2009).
For a careful evaluation of generic limits, which are best drawn broadly given the
extensive generic polyphyly and highly homoplasious "generic" characters, especially
within Cyclosorus s.l., see He and Zhang (2012).

[Woodsiaceae [Athyriaceae [Blechnaceae + Onocleaceae]]]: ?

WOODSIACEAE Herter

Plant epipetric; rhizomes suberect; petiole bases persist; circumendodermal band

surrounding leaf trace 0; indusium basal, of many scale-like or filamentous segments,
receptacle raised; n = 33, 38, 39, 41. 1/35. Mostly montane, northern hemisphere.

[Athyriaceae [Blechnaceae + Onocleaceae]]: ?

Woodsia may be sister to this whole clade (Schuettpelz & Pryer 2009: Fig. S1).
ATHYRIACEAE Alston

Mature frond with abundant anthocyanin; margins entire, petiole base swollen,
(starch-containing), persistent [= trophopod]; corniculae/scales at adaxial junction of
pinna costa with rachis; indusia opening to face away from a single vein [either two linear
back to back sori, or J-shaped indusium of a single sorus wrapped around the vein]; n =
40, 41. 5/600: Diplazium (400). Mostly, terrestrial, understory.
Age. Crown-group Athyriaceae are around 78.4 m.y.o. (Schuettpelz & Pryer 2009).
Wei et al. (2013) evaluated relationships within Diplazium and found i.a. that
species previously assigned to Allantodia in particular were scattered through the tree;
they circumscribed Diplazium broadly and provided an infrageneric classification for it.
For divergence dates and biogeography in Diplazium, perhaps a member of the
boreotropical flora in the Eocene, see Wei et al. (2015).
[Blechnaceae + Onocleaceae]]: fronds dimorphic [fertile and sterile].
BLECHNACEAE Newman

Young leaves reddish; leaf traces several, from V-shaped bundle; petiole abaxially with
three to many round vascular bundles arranged in an arc; (fronds monomorphic),
veins forming narrow areoles near the costa; sori linear, on subcostal commissural vein,
indusia opening towards costa; perine winged; n = 34 [27, 28, 31-37, 40]. 4/200.
Cosmopolitan

Age. Crown-group Blechnaceae are around 59.8 m.y.o. (Schuettpelz & Pryer 2009).
For relationships in Blechnaceae, with Blechnum circumscribed rather broadly, see
Perrie et al. (2014).
ONOCLEACEAE Pichi-Sermolli

Circumendodermal band surrounding leaf trace 0; petiole basally swollen, vascular

bundles uniting distally into a gutter shape; trophopods +; fronds dimorphic; sori
enclosed by reflexed lamina margins; indusium deltate; spores chlorophyllous; n =
37, 39, 40. 4/5. Northern Hemisphere.
Age. Onoclea sensibilis is known fossil from Palaeocene North America 62-58 m.y.a.,
the fossils being remarkably similar to extant individuals (Rothwell & Stockey 1991).

MONILOPHYTA / pakis S.L.

Akar dengan sel apikal tunggal, yang berasal dari Pericycle, akar lateral dari
endodermis; batang dengan awal apikal (s), kepadatan plasmodesmatal di
seluruh SAM 19-56 [berarti] / m2 [jaringan mitokondria garis keturunan
khusus]; batang dengan hypodermal dan band luar-kortikal serat [= stereome];
siphonostele amphiphloic +, diskontinuitas dalam prasasti di t.s. disebabkan oleh
kesenjangan pakis; protoxylem dibatasi untuk lobus tengah xilem untai
[memberikan penampilan manik-manik, maka monilophytes], xilem mesarch,
tracheids dengan lubang-lubang scalariform, G-jenis tracheids di protoxylem;
serat floem langka; batang endodermis dan Pericycle +; daun megaphyllous
[iklan / abaxial simetri berkembang pertama, kemudian determinancy],
pengembangan acropetal; tangkai daun dengan jejak daun beberapa yang
datang dari bundel berbentuk U; pelepah urat tidak anastomosing; sporangia
dikelompokkan dalam sori, sporangium tangkai 6 <sel di seluruh, dinding tebal
dua sel, dehiscence oleh exothecium, tapetum amoeboid, spora / sporangium
1000 <, putih, bulat-tetrahedral, dengan orbicules, pembangunan dinding
sentrifugal, exospore 3-lapis, pseudoendospore +; gametofit thalloid; dinding
antheridium 5 sel tebal, gamet jantan dengan 30-150 silia, dengan berbagai
plastida dan mitokondria; akar lateral yang berkenaan dengan sumbu
longitudinal dari embrio [tanaman homorhizic]; x = ca 121, ukuran genom [1C] =
ca 14,3 pg [rata-rata], gen rps4 kloroplas dengan penyisipan sembilan
nukleotida, inversi LSC dari Lthn-GCC untuk trnT-GGU; kehilangan satu kelompok
II intron mitokondria.
Usia. Magallon et al. (2013) memperkirakan usia mahkota-kelompok sekitar
(404-) 394.3-389.9 (-382) m.y. untuk clade ini, ca 360 dan ca 364 m.y. adalah
usia di Schuettpelz dan Pryer (2007) dan Y-L. Qiu et al. (2007) masing-masing,
(390.7-) 368,5 (-354) m.y. di Zhong et al. (2014b) dan sekitar 330 m.y. di
Villarreal dan Renner (2014); lihat juga P. Soltis et al. (2002) untuk saran.
Evolusi. Divergence & Distribution. Schuettpelz dan Pryer (2009, esp. Tabel 2, 3
di Tambahan) dan Rothfels et al. (2015b: Lampiran S4) memberikan kencan
ekstensif kali divergensi di monilophytes, dan untuk usia lebih dari clades
monilophyte utama melihat Y.-L. Qiu et al. (2007) dan Schneider et al. (2004a).
Schuettpelz dan Pryer (2009) juga daftar sejumlah catatan fosil (lihat juga
Rothwell & Stockey 2008).
Ada beberapa radiasi pakis leptosporangiate homosporous, yang pertama di
Paleozoikum, sehingga menimbulkan garis keturunan yang telah sejak punah,
dan lagi di Jurassic dan Cretaceous (Rothwell & Stockey 2008). keragaman pakis
umum menurun (bersama dengan yang dari sikas) melalui Kapur (Wing &
Boucher 1998). Pakis tampaknya telah sementara mendominasi setidaknya
secara lokal setelah bolide dampak / letusan Deccan Traps akhir-Kapur
(Schneider et al. 2004a). The eusporangiate Marattia dan Angiopteris dan
leptosporangiate kerajaan dan pohon pakis hampir fosil hidup dan menunjukkan
sedikit molekul dan bahkan evolusi morfologi (P. Soltis et al, 2002;. Zhong et al
2014b; Rothfels et al 2015b;.. Bomfleur et al 2014a, cf Schneider et al. 2015).
Beberapa 98% dari pakis hidup yang pakis leptosporangiate, dan radiasi utama
mereka mungkin memiliki telah setelah diversifikasi angiosperma pada akhir
Cretaceous-awal Caenozoic; awalnya itu adalah radiasi sebagian besar terestrial
(Lovis 1977; Schneider et al 2004a;. Rothwell & Stockey 2008; Schuettpelz &

Pryer 2009; H.-M. Liu et al 2014.). Lebih dari 70% dari pakis yang eupolypods,
dan sekitar sepertiga dari ini bersifat epifit, amd ini membuat ca 10% dari semua
epifit vaskular. Diversifikasi pakis epifit khususnya terjadi selama Paleogen,
mungkin terkait dengan Palaeocene-Eosen maksimum termal (Schneider et al
2004a, b; Schuettpelz 2007; esp Schuettpelz & Pryer 2009:.. Tabel Tambahan 2,
3; Watkins et al 2010. ; cf Dubuisson et al 2009, sebagian).. Pengecualian adalah
Trichomanes dan kerabat (tapi tidak Hymenophyllum dan kerabatnya, yang,
melihat Dubuisson et al 2009.) Yang terdiversifikasi di awal Cretaceous; mereka
umumnya epifit pada pohon pakis, yang sendiri telah dimulai diversifikasi di
Jurassic yang (Schuettpelz 2007; juga Schuettpelz & Pryer 2009; Rothwell &
Stockey 2008 untuk radiasi awal pakis leptosporangiate).
Sebuah hirarki sementara karakter yang diambil dari Smith et al. (2006, 2008),
Pryer et al. (1996) dan Rothfels et al. (2012b) diberikan di bawah ini. Untuk
menusuk pertama di apomorphies untuk anggota clade polypod II, melihat
Sundue dan Rothfels (2013); saran-saran mereka sebagian besar diikuti di sini;
masalah dengan pengambilan sampel, karakter negara batas, dan apomorphies
yang berbeda yang dihasilkan oleh metode yang berbeda dari optimasi karakter
diperhitungkan sejauh mungkin. Hanya synapomorphies ambigu (Sundue &
Rothfels 2013) ditandai bawah. Kaplan (.. 1997, vol 3: chap 19) diringkas
morfologi dinding sporangium dari monilophyta; anulus merupakan apomorphy
dari Polypodiopsida, pakis leptosporangiate. Untuk apomorphies lain yang
mungkin, lihat misalnya Schneider et al. (2009). Lebih banyak fitur perlu
ditambahkan, misalnya jika daun megaphyllous paling pakis dan mereka bibit
tanaman telah berevolusi secara independen, yang nampaknya (lihat di atas).
Memahami evolusi tubuh tanaman tampaknya sangat unfern-seperti Psilotales
dan Psilotum sendiri, sampai cukup baru-baru dianggap mungkin yang paling
primitif yang masih ada tanaman vaskular - ketika saya diajarkan tentang
Psilotum, itu dibandingkan dengan rhyniophytes Paleozoikum - masih hadiah
kesulitan. Kaplan (1997, vol 3:.. Chap 10; Siegert 1973 dan referensi) diringkas
literatur, mencatat bahwa masalah ini adalah karena morfologi Psilotum
khususnya dibandingkan dengan tanaman fosil ini, sekarang dianggap sama
sekali tidak berhubungan. Saat ia mencatat, daun muda dari Psilotum memiliki
fitur daun (lihat juga Kaplan 1977). Demikian pula, daun Equisetum mungkin
sekunder sederhana. Sehingga relatif Sphenophyllum fosil memiliki daun yang
lebih besar dengan venation dikotomis; xilem bifacial telah ditemukan dalam
genus ini (Kenrick & Derek 1997; Doyle 2013 dan referensi).
Ekologi & Physiology. Fisiologi evolusi dari pakis yang membayar pemeriksaan.
The tracheids cenderung panjang dan lebar, dan scalariform perforasi piring
dapat memperpanjang panjang sel. Akibatnya, transportasi air dapat relatif
efisien; konifer, kelompok lain tanpa kapal dan yang juga memiliki transportasi
air cukup efisien setidaknya memiliki penebalan sekunder dan lubang
berbatasan (Pittermann et al. 2011). kekayaan pakis di Australia berkorelasi
dengan curah hujan, dan tergantung pada apa aspek keanekaragaman
ditekankan, curah hujan mungkin terkait dengan ketetapan sifatnya, dll,
bryophytes, bukan tidak berbunga tanaman, menunjukkan pola yang agak mirip
(Nagalingum et al. 2015).
pakis epifit terkonsentrasi di hymenophyllaceae, Polypodiaceae, pteridaceae (mis
vittaria) dan dryopteridaceae (mis Elaphoglossum) (Schuettpelz & Pryer 2009;
Kato & Tsutsumi 2013; lihat juga di atas). The gametophytes epifit ini sering tali
berbentuk dan berumur panjang, dan terutama tahan terhadap dessication;

sebagai Watkins et al. (. 2007: p 716) mengamati, "gametophytes pakis yang,

untuk semua maksud dan tujuan, bryophytes", dan spesies epifit mereka yang
diteliti yang paling tahan (lihat juga Nayar & Kaur 1971: survei keanekaragaman
gametofit; Dassler & Farrar 2001 ; Farrar et al 2008;. Watkins & Cardels 2012;
Rothfels & Schuettpelz 2014). Gemmae pada gametophytes ini (mis Pryer et al.
2016) lebih lanjut akan meningkatkan umur panjang mereka. Pakis menunjukkan
berbagai adaptasi lain untuk habitat epifit, termasuk CAM fotosintesis (Watkins &
Cardels 2012) dan produksi Gemmae (Farrar 1974), dan mikoriza kurang sering
(mis B. Wang & Qiu 2006; Kato & Tsutsumi 2013); nutrisi cenderung rendah di
habitat epifit. Testo dan Sundue (2014) menemukan bahwa kedua spesies darat
dan hemiepiphytes (di mana akar dari tanaman memanjat membangun koneksi
dengan tanah) berasal dalam clade dari epifit Polypodiaceae, sedangkan J. Clark
et al. (2016) menyarankan bahwa mungkin ada hubungan antara kebiasaan
epifit dan peningkatan ukuran genom di eupolypods.
Kawai et al. (2003) menemukan khas chimaeric merah / jauh lampu merah
fotoreseptor (phy 3, = neochrome) di mana fitokrom merah-sensing dan
phototropin biru-sensing yang menyatu menjadi satu molekul (F.-W. Li et al.
2014) di beberapa Polypodiales, mungkin membantu dalam tanggapan
phototropic dalam kondisi teduh di mana pakis telah terdiversifikasi; neochrome
sangat sensitif terhadap cahaya putih (Wada 2008). Neochrome tampaknya telah
diakuisisi oleh pakis polypod dari hornworts melalui transfer gen lateral sekitar
179 mya, yang, mungkin tidak begitu kebetulan, adalah tentang usia
Polypodiales (Schneider et al. 2004a), dan mungkin bahkan telah mengakuisisi
lebih dari sekali ( F.-W. Li et al. 2014).
Pada tumbuhan biji, stomata terbuka di respon baik lampu merah dan biru.
Stomata dari mereka beberapa pakis diperiksa, semua leptosporangiates (Pteris,
Adiantum, Asplenium dan Nephrolepis), kurang respon pembukaan cahaya
tertentu biru, meskipun elemen dari respon, termasuk phototropin relevan, hak
kinase protein reseptor biru, yang hadir (Doi et al. 2006). Dalam lingkungan
understory berbayang bahwa banyak pakis lebih, cahaya biru (dan merah) telah
istimewa diserap oleh kanopi di atas; penutupan stomata dimediasi oleh lampu
merah. Selanjutnya, pakis tidak memiliki daun di mana pembuluh darah adalah
jarak yang sama dari epidermis karena mereka satu sama lain; mereka tidak
hidrolik dioptimalkan. Namun, fotosintesis dapat melanjutkan terbaik di tingkat
rendah cahaya menyebar dan optimasi hidrolik ini mungkin tidak begitu penting
(Zwieniecki & Boyce 2014). (Dalam tumbuhan berbiji efektivitas respon putih /
biru + merah> biru> merah> lampu hijau [Willmer & Fricker 1996] dan dalam
banyak tanaman berbunga daun hidrolik dioptimalkan [Zwieniecki & Boyce
2014].)
Pemupukan & Spore Penyebaran. Distribusi tanaman sporophytic dan
gametophytic dari spesies satu pakis mungkin sangat berbeda, terutama jika
gametofit adalah selain hati-berbentuk. gametophytes tali berbentuk atau
filamen dapat hidup untuk waktu yang sangat lama dan / atau menghasilkan
Gemmae, dan sehingga mereka dapat bertahan dalam situs yang ratusan mil
dari sporofit terdekat (mis Farrar 1967; Ebihara et al 2013.). Dengan munculnya
kemampuan untuk mengidentifikasi gametophytes langsung oleh sequencing
molekul, daripada menunggu mereka untuk menghasilkan sporofit, fenomena ini
berubah menjadi sangat umum (Ebihara et al. 2013), dan memiliki implikasi
yang menarik bagi evolusi pakis (lihat misalnya Ebihara et al. 2009).
Noblin et al. (2012) menjelaskan mekanisme bagaimana fungsi anulus di

sporangium dehiscence.
Tanaman-Animal Interaksi. Secara keseluruhan, herbivora di pakis adalah hampir
sama dengan rata-rata untuk angiosperma (Turcotte et al 2014:. Lihat
peringatan).
Bakteri / Asosiasi jamur. Untuk mikoriza di pakis, melihat Lehnert et al. (2010 dan
referensi). Asosiasi mikoriza tidak dikenal di Equisetum (Baca et al. 2000).
Gen & Genom. Untuk evolusi monilophyte kloroplas genom, melihat Karol et al.
(2010), Grewe et al. (2013) dan J. Clark et al. (2016). Dalam Polypodiopsida,
beberapa inversi di kloroplas terbalik berulang mungkin tingkat tinggi
synapomorphies (Gao et al. 2009). Grammitidaceae s. str. khususnya (termasuk
dalam Polypodiaceae) memiliki spora hijau dan dipercepat plastid genom evolusi,
korelasi ditemukan di tempat lain di pakis, meskipun tidak 100% (Schneider et
al. 2004b), memang, spora yang kurang jelas mengandung kloroplas yang cukup
luas (Sundue et al. 2011).
Pakis dicatat untuk insiden yang tinggi poliploidi, hampir 1/3 (31%) dari semua
peristiwa spesiasi yang disertai dengan poliploidi (Wood et al. 2009), dan nomor
dasar kromosom juga biasanya sangat tinggi. Telah hibridisasi baru-baru ini
antara clades (Cystopteris, Gymnocarpium) yang menyimpang diperkirakan
(76.2-) 57,9 (-40,2) m.y.a. (. Rothfels et al 2015a) sementara ada hibridisasi di
Osmunda antara clades yang mungkin telah terpisah selama empat kali selama
(Bomfleur et al 2014b;.. Grimm et al 2015).
Untuk informasi tentang ukuran genom, melihat Obermayer et al. (2002) dan J.
Clark et al. (2016); terutama genom besar terjadi pada Psilotales dan beberapa
Ophioglossales, dan dalam beberapa polypods.
Kimia, Morfologi, dll Komposisi xyloglucan dari dinding sel primer bervariasi
secara substansial, bahwa dari Equisetum menjadi sangat khas (Hsieh & Harris
2012). Untuk polisakarida dinding sel, lihat juga Silva et al. (2011).
morfologi sporophyte dasar telah digariskan oleh Kaplan (1997, vol 2:.. chap 11,
18, vol 3:.. chap 19, 2001). Untuk organisasi dari meristem apikal, melihat
Amrbose dan Vasco (2015). batang memiliki siphonostele sebuah, protoxylem
yang dibatasi untuk lobus untai xilem pusat, maka membawa ke pikiran kalung
(pengembangan xilem adalah mesarch, meskipun terutama variabel dalam clade
ophioglossum / Psilotum). protoxylem yang digambarkan memiliki G-jenis
tracheids (Edwards 1993). Hernndez-Hernndez et al. (2012) membahas
distribusi band circumendodermal, tannin yang mengandung sel-sel yang
mengelilingi jejak daun petiolar yang memiliki orgin umum dengan endodermis;
mereka juga detail distribusi sejumlah fitur vegetatif / kebiasaan lainnya. Vasco
et al. (2013) meringkas pakis daun morfologi dan pengembangan, mencatat
sejumlah fitur tembak-seperti. Davies (1991) diringkas informasi tentang
aerophore distribusi di pakis. Informasi pada dinding sel horizontal dalam
perkembangan embrio awal pakis diberikan oleh Philipson (1990) tampaknya
menjadi salah - contoh harus vertikal?
Untuk puncak batang pakis, lihat di atas.
Takahashi et al. (2009, 2014 dan referensi) menjelaskan pengembangan
gametofit di pakis. Mereka mencatat bahwa wilayah apikal mengkonversi ke

meristem multiseluler, yang dapat membagi - percabangan dikotomis - jika

pembelahan sel di tengah meristem berhenti; bercabang, gametophytes tali
berbentuk pakis epifit hanya merupakan varian ekstrim morfologi ini. Archegonia
berkembang hanya setelah pembentukan meristem multiseluler. Untuk rincian
morfologi gamet jantan dan gerakan, dll, lihat misalnya Renzaglia et al. (2000b,
2002) dan Schneider et al. (2002).
Untuk informasi tentang pteridophytes pada umumnya (ini memiliki sering - dan
masih mungkin - termasuk lycophyte), lihat juga Kato (2005) dan Ranker dan
Haufler (2008). Untuk informasi umum lainnya, lihat Raven dan Edwards (2001),
untuk anatomi perbandingan, lihat Ogura (1972), untuk kapal, melihat Sen dan
Mukhopadhyay (2014 dan referensi), untuk venation, melihat Wagner (1979) dan
Boyce (2005b), untuk rincian morfologi stele dan evolusi, melihat Beck et al.
(1982), dan untuk sporofit muda, dll, lihat Johnson dan Renzaglia (2009 dan
referensi).
Filogeni. The batasan dari clade ini hanya baru-baru telah menjadi jelas. Ini
termasuk sangat mendukung [Psilotum + ophioglossum] clade (Tmesipteris
adalah adik untuk Psilotum) mungkin saudara semua pakis lain, sebagai data
kloroplas telah secara luas cenderung menyarankan (Rothfels et al. 2015b untuk
referensi). Wickett et al. (2014) memperoleh [Marattiales + Psilotales] clade adik
leptosporangiate pakis, tapi ini mungkin menjadi masalah sampling.
Dimasukkannya morfologi sendiri atau dalam kombinasi dengan data molekuler
juga mempengaruhi hubungan terdeteksi (Wikstrom & Pryer 2005 dan referensi);
lihat juga Grand et al. (2013) untuk berbagai analisis morfologi. Posisi Equisetum
tidak pasti.
1. Mungkin saudara Angiopteris, dll (walaupun dukungan hanya moderat), clade
gabungan pada gilirannya menjadi saudara pakis yang tersisa (misalnya Pryer et
el 2001a, 2004a;. Wikstrom & Pryer 2005; Qiu et al 2007;. Ebihara et al 2011;. cf
di bagian Serigala et al 1998)..
2. Mungkin saudara semua pakis lain, seperti dalam analisis rps4, dan juga
menganalisis 4- dan 5-gen, dua terakhir dengan dukungan yang kuat
(Schuettpelz et al. 2006), analisis beberapa gen plastid (Rai & Graham 2010),
dan dalam filogeni matK (Kuo et al. 2011). Knie et al. (2015) juga menemukan
dukungan yang baik untuk hubungan [Equisetum [[Psilotum + ophioglossum] +
The Rest]], seperti halnya Rothfels et al. (2015b) dalam analisis gen nuklir
mereka. Menariknya, spora dinding ultrastruktur dari calamites, pabrik
equisetaceous punah, tidak begitu berbeda dari ophioglossaceae dan pakis
lainnya (Lugardon & Brousmiche-Delcambre 1994; Grauvogel-Stamm & Lugardon
2009). Equisetum memiliki atp1 intron mitokondria, baik sekunder (dan sejajar)
kehilangan atau tidak adanya plesiomorphic, tergantung pada topologi dari
seluruh kelompok (Wikstrom dan Pryer 2005: melihat hirarki karakter di bawah
ini).
3. Schneider et al. (2009) mencatat potensi apomorphies morfologi seperti helai
daun sederhana dan batang dengan kedua radial dan dorsiventral simetri (=
tegak ditambah merayap batang ...) menunjukkan clade [Psilotales +
Equisetales], konsisten dengan beberapa perubahan struktural dalam genom
kloroplas (Grewe . et al 2013; lihat juga beberapa analisis di Karol et al 2010;
juga Serigala & Karol 2012; Ruhfel et al 2014; J.-M. Lu et al 2015:.. genom
kloroplas)..

reorganisasi ini monilophytes kadang-kadang dikritik (Rothwell & Nixon 2006),

tetapi tidak jelas bagaimana memberatkan kritik tersebut. Karena evaluasi
"dukungan" nilai untuk topologi tertentu merupakan bagian integral pendekatan
yang digunakan dalam halaman-halaman ini, keputusan untuk mengecualikan
nilai-nilai tersebut oleh mereka penulis membuat pekerjaan mereka sulit (bagi
saya, setidaknya) untuk menafsirkan. Memang, dalam beberapa analisis cladistic
morfologi (mis Bremer 1985; Stevenson & Loconte 1996; Rothwell 1999: fosil
termasuk atau tidak) Psilotum keluar sebagai saudara semua tumbuhan vaskular
lainnya. Namun, beberapa analisis morfologi (Schneider et al. 2009) melakukan
tempat Psilotum dengan monilophytes lain, bahkan jika analisis yang sama juga
menempatkan tanaman berbunga dalam kelompok paraphyletic dari
gymnosperma yang masih ada.
Klasifikasi. Smith et al. (2006, 2008) mengusulkan reklasifikasi berbasis filogeni
dari pakis, dan mereka juga mencakup sastra, sinonim ordinal dan keluarga, dan
daftar yang diterima genera dan beberapa sinonim besar; Prelli (2010)
memberikan bagus akun pakis Eropa. Namun, penyesuaian klasifikasi ini sedang
dibuat sebagai rincian dari filogeni menjadi lebih baik dipahami (Schuettpelz &
Pryer 2007, 2008; Kuo et al 2011; Rothfels et al 2012b:.. Reklasifikasi eupolypods
II). Sebuah urutan linier dari keluarga dan genera (Christenhusz et al. 2011a)
sekarang tanggal, tetapi baru-baru Christenhusz dan Chase (2014) telah
mengusulkan klasifikasi lain. Ada beberapa perbedaan antara hierarki di bawah
ini dan yang digunakan oleh mis Rothfels et al. (2015b), tetapi mereka akan
dibersihkan sebagai hal-hal (mudah-mudahan) menetap.
Hubungan sebelumnya. Psilotum dan Equisetum sebelumnya dianggap mewakili
garis keturunan independen, dengan Psilotum dan kerabat dianggap paling
primitif yang hidup tumbuhan vaskular, dan yang terakhir memang terlihat
dangkal mirip dengan beberapa fosil awal. hubungan mereka dengan pakis,
sekarang sangat sebagian besar diterima, tak terduga (tapi lihat Kenrick & Derek
1997). Meskipun Bierhorst (1968, lihat juga 1977) dibandingkan Psilotum dengan
pakis yang masih ada Stromatopteris dan menemukan beberapa kesamaan
morfologi, sebagian besar telah berubah menjadi paralelisme dan keduanya
tidak sama sekali dekat.
EQUISETOPSIDA / [Equisetales [Psilotales + Ophioglossales]]: tanaman dengan
tegak dan merayap batang; ? Vernation; tapetum plasmodial; embrio exoscopic,
suspensor 0; gen rps16 kloroplas dan rps12i346 intron hilang.
Usia. Clade yang berisi Equisetum mungkin telah terpisah dari monilophytes
lainnya sejak Permian, ca 250 + m.y.a. (Stanich et al 2009.); Zhong et al. (2014b)
berpikir bahwa clade ini (370.3-) 296,2 (-189,9) m.y. tua.
EQUISETALES Berchtold & Presl
EQUISETACEAE de Candolle
Tanaman dengan tegak dan merayap batang; akar triarch, dengan trakeid pusat
besar; dinding sel juga dengan (1-> 3), (1-> 4) -sS-D-MLGs [Glucans-Linkage
Mixed], akumulasi Si02 umum; batang dengan meristem kabisat [di dasar daun
kelopak], bergerigi, fotosintesis, protoxylem mesarch, dengan kanal pusat;
kekosongan protoxylem berkembang; bundel daun vaskular amphicribral; cabang
whorled, anggota whorls bolak pada setiap node; daun kecil, sederhana, 1berurat, whorled, basally bawaan, tidak fotosintetik; sporangiospora peltate,
dikumpulkan ke dalam Strobilus sebuah; dinding sel sporangial dengan

thickenings sekunder heliks; tapetum plasmodial; spora dengan aperture

melingkar [hilate], abapertural obturator +, hijau, dinding dengan silika, elaters
4-6 / spora, spatulate, spiral-melingkar; embrio exoscopic, pesawat dari variabel
pembelahan sel pertama, suspensor 0; n = 108; mitokondria atp1 intron 0.
1/15. Dunia-lebar, bukan Antipodes.
Usia. spesies yang masih ada dari Equisetum tampaknya telah dipisahkan dalam
Caenozoic (77.5-) 64,8 (-52,1) m.y.a. (Des Marais et al. 2003, namun c.f. Stanich
et al. 2009).
Fosil dengan banyak apomorphies kelompok mahkota Equisetum dikenal dari
deposito Jurassic Atas dari Patagonia sekitar 150 m.y. atau lebih tua (Channing et
al 2011;. lihat juga Stanich et al 2009.); untuk spora Equisetum-seperti masih
lebih tua - tetapi dengan trilete tanda - dan elaters terkait dari Trias Tengah,
melihat Schwendemann et al. (2010).
Evolusi. Divergence & Distribution. Perubahan spora morfologi dari jenis
calamites ke pada pandangan pertama spora trilete sangat berbeda dari
Equisetum yang meyakinkan ditunjukkan oleh Grauvogel-Stamm dan Lugardon
(2009). fosil lainnya ditempatkan di daerah ini termasuk Sphenophyllum, yang
memiliki megaphylls, dan ini menunjukkan bahwa daun kecil Equisetum dapat
dikurangi bukannya microphylls. Akar dari fosil juga berbeda dalam morfologi
dan dapat polyarch.
Ekologi & Physiology. Equistetum cenderung tumbuh di habitat ekologis agak
stressfull, termasuk mata air panas (al Channing et 2011 dan sastra;. Husby
2012). Ada bertekanan aliran gas, yang disebut ventilasi konvektif, dari batang
ke dalam rimpang di beberapa spesies yang masih ada dari Equisetum dan
mungkin di kerabat calamitalean fosil mereka dari Karbon. Oksigen bergerak
melalui ruang udara yang saling berhubungan dalam rimpang, sehingga
memungkinkan mereka untuk menembus dalam ke substrat anoxic umumnya
disukai oleh kelompok ini (Armstrong & Armstrong 2009). Namun, mereka
spesies Equisetum yang tidak memiliki ruang udara yang saling berhubungan dan ini disertai dengan perubahan posisi dari endodermis aliran-tahan di batang
- tidak memiliki ventilasi konvektif (Armstrong & Armstrong 2011).
Biologi reproduksi. Dibantu oleh elaters mereka, spora Equisetum dapat baik
melompat hingga 1 cm di udara saat kering, atau bergerak dengan gerakan
pendek random walk-jenis sepanjang tanah (Marmottant et al. 2013).
Gen & Genom. Untuk duplikasi genom di Equisetum tanggal untuk (112.5-) 92,4
(-75,2) - atau mungkin hanya 70-50 - m.y.a., lihat Vanneste et al. (2015).
Kimia, Morfologi, dll Untuk (1-> 3,1-> 4) -sS-d-glukan, lihat Fry et al. (2008) dan
Xue dan Fry (2012), dan untuk konsentrasi Si, dll, lihat Husby (2012).
Filogeni. Equisetum bogotense adalah adik ke seluruh genus (Guillon 2007).
[[Psilotales + Ophioglossales] [Marattiopsida + Polypodiopsida]]: (spora aperture
proksimal dan monolete); INDEL di mitokondria Rpl2 wilayah coding.
Usia. Rothfels et al. (2015b) disarankan usia (390.4-) 351,7 (-309,7) m.y. untuk
node ini.

[Psilotales + Ophioglossales]: tanaman dengan tegak dan merayap batang;

batang variabel pengembangan protoxylem; embrio exoscopic, suspensor 0;
gametofit bawah tanah, aksial, non-fotosintetik, mikoriza; Ukuran genom nuklir
32,7-64,8 pg.
Usia. Magallon et al. (2013) memperkirakan usia sekitar 275,6 m.y. untuk clade
ini, dan (316-) 306 (-267) m.y. adalah usia di Y. L. Qiu et al. (2007), ca 214,5 m.y.
di Laenen et al. (2014) dan (317.1-) 250,5 (-141,8) m.y. di Rothfels et al. (2015b);
lihat juga P. Soltis et al. 2002) untuk perkiraan.
Gen & Genom. Kedua Psilotum dan ophioglossum memiliki genom besar dengan
(Bennett & Leitch 2005; J. Clark et al 2016.).
PSILOTALES Prantl
Psilotaceae J. W. Griffith & Henfrey
epifit; akar 0; ? Bundel daun pembuluh darah; daun kecil, sederhana, (lateral rata
- Tmesipteris), vena 1 atau 0; sporangia 2-3, membentuk synangium; tapetum
kelenjar-amoeboid; spora berbentuk ginjal, monolete; gametofit dengan rhizoids
septate; (Sel transfer dalam sporophyte hanya - Psilotum), n = 52, 208, ukuran
genom [1C] = ca 72,5 pg. 2/12.
Usia. Zhong et al. (2014b) memperkirakan usia (147.1-) 72,3 (-14,7) m.y. dan
Rothfels et al. (2015b) usia (142.5-) 78,9 (-28,5) m.y.a. untuk clade ini.
OPHIOGLOSSSALES Tautan
ophioglossaceae Martinov
Root dengan 2-5 tiang protoxylem; akar rambut 0; gabus pertengahan kortikal;
vaskular kambium +; batang sympodial prasasti; tracheids dengan melingkar
berbatasan lubang; daun vaskular bundel agunan; (Tunas ketiak +); daun
majemuk untuk sederhana, 1 diproduksi / tahun, venation retikular, dengan urat
diarahkan secara internal, vernation mengangguk, basis selubung, stipula +,
tipis; satu atau lebih sporophores terkait dengan setiap tropophore; (Gametofit
dengan rhizoids septate); (Embrio endoskopik; dinding sel pertama dari zigot
vertikal); n = (44) 45 (46 ... 720), ukuran genom [1C] = ca 28,35 pg. 4/80.
Usia. Rothfels et al. (2015b) disarankan usia (249.6-) 161,7 (-74) m.y. untuk
mahkota-kelompok ophioglossaceae.
mikoriza Glomeromycote di ophioglossum berhubungan dengan gametofit
echlorophyllous dan tahap sporophytic bawah tanah, dan juga sporophyte
photosynthesising (Field et al. 2015a). Takahashi dan Kato (1988)
menggambarkan perkembangan meristem lateral dalam keluarga.
reticulatum ophioglossum, di n = 720, memiliki jumlah kromosom tertinggi
setiap tanaman.
Lihat Hauk et al. (2003) untuk filogeni, Mankyua tidak termasuk, juga Shinohara
et al. (2013), Mankyua termasuk, tapi posisi tidak stabil - adik untuk seluruh
keluarga (juga dalam analisis bersama), atau untuk ophioglossum s. str ..
[Marattiopsida + Polypodiopsida]: bundel daun vaskular amphicribral; senyawa

daun palem, vernation circinate; sisik +; sporangia abaxial; gametofit hijau,

surficial; menembak berkembang menuju leher archegonial [dari sel hypobasal,
endoskopi]; nuklir genom 3,5-14,0 pg [disini?]; adjacencies gen berubah di
perbatasan IR kloroplas; mitokondria gain atp1i361g2 intron.
Usia. Zhong et al. (2014b) disarankan usia (378.8-) 336,7 (-291,5) dan Rothfels
et al. (2015b) usia (364.1-) 329 (-289,2) m.y. untuk clade ini.
P. Soltis et al. (2002) menawarkan berbagai saran untuk simpul umur dalam
clade ini.
MARATTIOPSIDA
Akar dengan beberapa tiang protoxylem; rambut akar septate [multiseluler?];
tangkai daun pembuluh darah polycyclic; stipula +, berdaging dan tepung.
MARATTIALES Tautan
Sinonim: Christenseniales Doweld
marattiaceae Kaulfman
Dictyostele +; kanal lendir +; rimpang dengan sisik; aerophores linear, dengan
lentisel; daun pulvinate, (selebaran dengan diarahkan internal retikular venation
- Christensenia); meiosis monoplastidic [semua}; spora bilateral atau ellipsoid,
monolete; sel Transfer 0; x = 40. 5/150: Danaea (50).
Untuk meiosis, melihat Brown dan Lemmon (2001).
Evolusi. Divergence & Distribution. Untuk beberapa komentar di biogeografi,
melihat Christenhusz dan Chase (2013). Tampaknya ada lambat-down dalam
tingkat evolusi dalam clade ini (Rothfels et al. 2015b dan referensi).
Filogeni. Untuk filogeni, lihat Murdock (2008a), juga Christenhusz et al. (2008);
yang Psaronius fosil tampaknya mengasosiasikan konsisten dengan Marattia (mis
Grand et al. 2013 dan referensi).
Kedua Marattia dan Angiopteris yang paraphyletic, tetapi mereka dapat dengan
mudah dibuat monofiletik (Murdock 2008b). Untuk Danaea, lihat Christenhusz
(2010).
pakis POLYPODIOPSIDA / leptosporangiate.
dinding sel primer miskin di mannans dan kaya tanin; akar dengan 2 kutub
protoxylem; xilem primer dengan scalariform berbatasan lubang; daun jejak
tunggal; aerophores linear, di kedua sisi tangkai daun, dengan stomata;
sporangium berasal dari divisi periklinal dari sel epidermis tunggal, dinding satulapis, tangkai 4-6 sel di seluruh [= leptosporangiate]; sporangium dengan
exothecium membentuk annulus, 64-800 spora / sporangium; antheridium
terkena; gametofit berbentuk hati [tingkat?]; embrio rawan [dinding sel pertama
dari zigot vertikal, sejajar dengan gravitasi], dengan kuadran / pembentukan
oktan, suspensor 0.
Usia. Magallon et al. (2013: dengan kendala sementara) diperkirakan usia sekitar
(267.8-) 252.7-251.4 (-246,1) m.y. untuk clade ini; (330) 323 (-310) m.y. adalah
usia di Y. L. Qiu et al. (2007), mungkin 350 m.y.a. di Schneider et al. (2004a), ca

299 m.y.a. di Schuettpelz dan Pryer (2009), dan sekitar 170 m.y.a. di Villarreal
dan Renner (2014) dan (327.8-) 301,3 (-271,5) m.y. di Rothfels et al. (2015b) semua mengatakan, penyebaran agak membingungkan usia.
Filogeni. Clade besar terdiri dari pakis leptosporangiate memiliki dukungan yang
sangat kuat (lihat juga Hasebe et al 1994, 1995, Pryer et al 1995;.. Serigala et al
1998;. Quandt et al 2004;. Schuettpelz et al 2006;. Rai & Graham 2010, dll).
Rothfels et al. (2015b) menekankan bahwa analisis mereka data nuklir secara
luas setuju dengan beberapa urutan plastid analisis.
Dalam leptosporangiates, Osmunda dan kerabat, sporangia yang memiliki
beberapa fitur eusporangiate, sangat didukung sebagai saudara sisanya. Ada
resolusi lebih lanjut substansial hubungan (mis Pryer et al 2004a, referensi b
dan; Schuettpelz & Pryer 2007; c.f. di bagian Kuo et al 2011:. Posisi
gleicheniaceae, lindsaeaceae, Nephrolepis [sebelumnya di lomariopsidaceae]
pasti.). Tidak jelas apakah atau tidak ada clade [Hymenophyllales +
gleicheniales] (Knie et al 2015 untuk sastra; Rothfels et al 2015b:. Gleicheniales
paraphyletic, tapi dengan sedikit dukungan.). J.-M. Lu et al. (2015: kloroplas
genom, tapi sampling) menemukan dipteridaceae dan Lygodiaceae menjadi
cabang berturut sepanjang batang leptosporangiate. Schizaeales dan paku air
(dukungan yang kuat) dan cyatheales (dukungan yang lemah) yang berturutturut saudara Polypodiales (mis Rothfels et al. 2015), dan Rai dan Graham (2010)
menyarankan varian tambahan. Untuk hubungan dengan schizaeales, melihat
Labiak et al. (2015 dan referensi).
Dalam Polypodiales, lindsaeaceae mungkin saudara semua orang lain (lihat juga
Rothfels et al 2015b.), Tetapi genera Cystodium, mantan dicksoniaceae, dan
Lonchitis dan Saccoloma, baik mantan Dennstaediaceae - yang terakhir sebagai
sebuah keluarga yang terpisah di bawah - juga di daerah ini (Lehtonen et al.
2012). Pteridaceae dan Dennstaediaceae baik didukung berturut taksa adik ke
eupolypods (Rothfels et al 2015b;. J.-M. Lu et al 2015.). Davalliaceae dan taksa
terkait adalah saudara pakis polygrammoid, dan keduanya mencakup sejumlah
epifit (evolusi mereka, melihat Tsutsumi & Kato 2006). Hubungan yang
disarankan oleh perubahan struktural dalam genom kloroplas (Wolf & Roper
2008;. Serigala et al 2010, 2011) konsisten dengan yang disarankan oleh urutan
analisis. Cystopteris dan kerabat membentuk clade yang mungkin adik ke clade
eupolypod II (Rothfels et al. 2009, esp. 2012a, 2013, 2015b).

OPHIOGLOSSSALES Tautan
ophioglossaceae Martinov
Root dengan 2-5 tiang protoxylem; akar rambut 0; gabus pertengahan kortikal;
vaskular kambium +; batang sympodial prasasti; tracheids dengan melingkar
berbatasan lubang; daun vaskular bundel agunan; (Tunas ketiak +); daun
majemuk untuk sederhana, 1 diproduksi / tahun, venation retikular, dengan urat
diarahkan secara internal, vernation mengangguk, basis selubung, stipula +,
tipis; satu atau lebih sporophores terkait dengan setiap tropophore; (Gametofit
dengan rhizoids septate); (Embrio endoskopik; dinding sel pertama dari zigot
vertikal); n = (44) 45 (46 ... 720), ukuran genom [1C] = ca 28,35 pg. 4/80.
Usia. Rothfels et al. (2015b) disarankan usia (249.6-) 161,7 (-74) m.y. untuk
mahkota-kelompok ophioglossaceae.
mikoriza Glomeromycote di ophioglossum berhubungan dengan gametofit
echlorophyllous dan tahap sporophytic bawah tanah, dan juga sporophyte
photosynthesising (Field et al. 2015a). Takahashi dan Kato (1988)
menggambarkan perkembangan meristem lateral dalam keluarga.
reticulatum ophioglossum, di n = 720, memiliki jumlah kromosom tertinggi
setiap tanaman.
Lihat Hauk et al. (2003) untuk filogeni, Mankyua tidak termasuk, juga Shinohara
et al. (2013), Mankyua termasuk, tapi posisi tidak stabil - adik untuk seluruh
keluarga (juga dalam analisis bersama), atau untuk ophioglossum s. str ..
[Marattiopsida + Polypodiopsida]: bundel daun vaskular amphicribral; senyawa
daun palem, vernation circinate; sisik +; sporangia abaxial; gametofit hijau,
surficial; menembak berkembang menuju leher archegonial [dari sel hypobasal,
endoskopi]; nuklir genom 3,5-14,0 pg [disini?]; adjacencies gen berubah di
perbatasan IR kloroplas; mitokondria gain atp1i361g2 intron.
Usia. Zhong et al. (2014b) disarankan usia (378.8-) 336,7 (-291,5) dan Rothfels
et al. (2015b) usia (364.1-) 329 (-289,2) m.y. untuk clade ini.
P. Soltis et al. (2002) menawarkan berbagai saran untuk simpul umur dalam
clade ini.
MARATTIOPSIDA
Akar dengan beberapa tiang protoxylem; rambut akar septate [multiseluler?];
tangkai daun pembuluh darah polycyclic; stipula +, berdaging dan tepung.
MARATTIALES Tautan
Sinonim: Christenseniales Doweld
marattiaceae Kaulfman
Dictyostele +; kanal lendir +; rimpang dengan sisik; aerophores linear, dengan
lentisel; daun pulvinate, (selebaran dengan diarahkan internal retikular venation
- Christensenia); meiosis monoplastidic [semua}; spora bilateral atau ellipsoid,
monolete; sel Transfer 0; x = 40. 5/150: Danaea (50).

Untuk meiosis, melihat Brown dan Lemmon (2001).

Evolusi. Divergence & Distribution. Untuk beberapa komentar di biogeografi,
melihat Christenhusz dan Chase (2013). Tampaknya ada lambat-down dalam
tingkat evolusi dalam clade ini (Rothfels et al. 2015b dan referensi).
Filogeni. Untuk filogeni, lihat Murdock (2008a), juga Christenhusz et al. (2008);
yang Psaronius fosil tampaknya mengasosiasikan konsisten dengan Marattia (mis
Grand et al. 2013 dan referensi).
Kedua Marattia dan Angiopteris yang paraphyletic, tetapi mereka dapat dengan
mudah dibuat monofiletik (Murdock 2008b). Untuk Danaea, lihat Christenhusz
(2010).
pakis POLYPODIOPSIDA / leptosporangiate.